Skip to main content

Full text of "The Journal of research on the lepidoptera"

See other formats










mm 






if 






i's|s:vi>3& 

Y*/ 






MJiw ''^J 

feXa 

%P 


iMti 

%^l 


^W X 


Mmm 











Qi 


'."^THE JOURNAL 
@NJ THE « 


@F RsESlA.^CHJ 


m. 


Established in 1 962 


Edited by WILLIAM HOVANITZ 


Volume 1 


1962 - 1963 


f 


Published at 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 


THE JOURNAL OF RESEARCH 


ON THE LEPIDOPTERA 


is sponsored by 


THE LEPIDOPTERA FOUNDATION 


CONTENTS 


Volume 1 

Number 1 

August, 1962 

Editorial 


1 


Variation in the silvering of Argynnis (Speyena) cdlippe in 
the interior mountain area of south central California . . . 

Oscar Elton Sette 3 


The effect of various food plants on survival and growth rate of 
Pieris .... William Hovanitz and Vincent C. S. Chang 21 

General characteristics of the movements of Vanessa cardui (L.) 

J. W. Tffden 43 

Three factors affecting larval choice of food plant 

William Hovanitz and Vincent C. S. Chang 51 

The generic, specific and lower category names of the Nearctic 
butterflies. Part 1 •— The genus Pieris . . Paddy McHenry 63 

The distribution of the species of the genus Pieris in North 
America William Hovanitz 73 

Did the caterpillar exterminate the giant reptile? .... 

S. E. Flanders 85 

Further evidence of the distribution of some boreal Lepidoptera 
in the Sierra Nevada Clyde Eriksen 89 

Argynnis and Speyer ia . William Hovanitz 94 


Volume 1 Number 2 January, 1963 

Composition and relative abundance in a temperate zone 
butterfly fauna . . . Thomas C. Emmel and John F. Emmel 97 

The Argynnis populations of the Sand Creek area, Klamath 
Co., Oregon, Part I .... J. W. Tilden 109 

Caterpillar Versus Dinosaur? . . . Theodore H. Eaton, Jr. 114 

Geographical distribution and variation of the Genus Argynnis 
I. Introduction 

II. Argynnis idalia William Hovanitz 117 

The relation of Pieris virginiensis Edw. to Pieris napi L. 

Species formation in Pieris? ..... Wiliam Hovanitz 124 

The male genitalia of some Co lias species . Bjorn Petersen 135 

The effect of hybridization of host-plant strains on growth rate 

and mortality of Pieris rapae . 

WiUiam Hovanitz and Vincent C. S. Chang 157 


Notice-editorial 


162 


Change of food plant preference by larvae of Pieris rapae 

controlled by strain selection, and the inheritance of this trait 

William Hovanitz and Vincent C. S. Chang 163 

Volume 1 Number 3 March, 1963 

Selection of allyl isothiocyanate by larvae of Pieris rapae and 

the inheritance of this trait 

William Hovanitz and Vincent C. S. Chang 169 

Biology of the Ceanothus stem-gall moth, Periploca ceanothiella 

(Cosens), with consideration of its control. J. Alex Munro 183 

Larval food-plant records for six western Papilios, .... 

John F. Emmel and ThtMnas C. Emmel 191 

CoUas philodice in Chiapa^ Mexico . . Thomas C. Emmel 194 

Notes on the early stages of two California geometrids 

John Adams Comstock 195 

Geographical distribution and variation of the genus Argynnis 

III. Argynnis diana William Hovanitz 201 

The generic, specific and lower category names of the nearctic 

butterflies. Part 2 — The Genus Colias . Paddy McHenry 209 

A standard method for mounting whole adult lepidoptera on 

slides utilizing polystyrene plastic . . Charles L. Hogue 223 

Volume 1 Number A May, 1963 


Techniques in the study of population structure in 
Philotes sonorensis 

Rudolf H. T. Mattoni and Marvin S. B. Seiger 237 
Early stages of a southern California Geometrid moth, 

Drepanulafrix hulsti hulsti (Dyar) John Adams Comstock 245 

The effeaiveness of different isothiocyanates on attraaing larvae 
of Pieris rapae 

William Hovanitz, Vincent C. S. Chang and Gerald Honch 249 

The origin of a sympatric species in Colias through the aid of 

natural hybridization William Hovanitz 261 

A method for breeding Pieris napi and Pieris bryoniae 

Bjorn Petersen 275 

An analysis of the North American species of the genus 

Callophrys J. w. Tilden 281 

THE LEPIDOPTERA FOUNDATION 301 

Notice 302 






Volume 1 

Number 1 

August, 1962 



Established in 1 962 
Published at 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

Edited by WILLIAM HOVANITZ 

WITH EMPHASIS ON ENVIRONMENTALLY AND GENE- 
TICALLY INDUCED VARIATION, population analysis, evolution, 
phylogenetic taxonomy, zoogeography, comparative morphology, ecol- 
ogy, geographical variation, speciation, physiology, etc. In short, 
quality tuork on any aspect of research on the Lepidoptera. 

THE PURPOSE OE THE JOURNAL is to combine in one source 
the work in this field for the aid of students of this group of insects 
in a way not at present available. The JOURNAL will attempt to 
publish primarily only critical and complete papers of an analytical 
nature, though there will be a limited section devoted to shorter 
papers and notes. 

RATES: $ 8.00 per volume, personal subscription. 

$12.00 per volume, institutional subscription. 

Benefactor subscriptions are encouraged. These may be 
any amount over those above. 

All amounts are in U.S. dollars, payable in the U.S.A. 

AUTHORS ARE REQUESTED to refer to the Journal as an 
example of the form to be used in preparing their manuscripts. Fifty 
separates will be supplied to authors free; reprints will be sold at 
printer’s rates if ordered at time galley proofs are returned. If proofs 
are not returned promptly, the editor reserves the right to withhold 
publication, or to proceed with no responsibility. All issues of the 
Journal will be copyrighted; in submitting a paper each author agrees 
that the material has not been published elsewhere. 

MANUSCRIPTS AND SUBSCRIPTIONS should be mailed to the 
address noted above. 

BENEFACTORS Individuals or institutions who by reason of 
their interest in the welfare or research on the Lepidoptera, and in 
its publication, who wish to contribute beyond the cost of a subscription, 
will be listed here. 


BIO-METAL ASSOCIATES 
THOMAS C. EMMEL 
ANONYMOUS DONOR 
ANONYMOUS DONOR 


1 ( 1 ): 1,1962 


Journal of Research on the Lepidoptera 

1140 W. Orange Grove Ave., Arcadia, California, U,S.A. 
Copyright 1962 


EDITORIAL 


Starting a new journal is not an easy matter, nor is it a matter free 
of controversy. This is especially the case where a new journal, as this 
one, is started without the support of an existing society. This journal 
is intended as a meeting ground for papers on all aspects of the biology 
of the Lepidoptera, moths and butterflies alike. Too many entomolog- 
ical journals don’t offer publication of articles of the length necessary 
to present sound data and Completeness of effort. The Lepidoptera are 
a group of insects second to none in the scope of our knowledge about 
them: work in this fieM has reached the higher levels of learning and a 
journal is required to keep pace with this knowledge at the same level. 

A journal with a wide scope is our aim — narrow in the taxonomic 
sense but wide in the biological sense. The journal will attempt to 
publish primarily only critical and complete papers of an analytical na- 
ture, though there will be a limited section devoted to shorter papers 
and notes. There will be emphasis on environmentally and genetically 
induced variation, population analysis, evolution, phylogenetic taxon- 
omy, zoo-geography, comparative morphology, ecology, geographical 
variation, speciation, physiology, etc. 

It has been very gratifying to receive so many subscriptions on the 
basis of merely a brochure; every effort will be made to keep the journal 
up to the standard expected of us. The response in so short a time has 
served to show those who have generously provided the finances that a 
journal of this type has been needed and will fill a desirable niche. 

We are happy to publish in this first issue a number of papers 
which the authors were able to get ready on "short notice". There is 
a preponderance on one taxonomic group, but with different points in 
mind. With this first issue out, we offer the pages of this journal to all 
with papers to publish on the Lepidoptera that fit the intent of the 
journal. A wider range of papers will come in future issues. 


1 




v:,.-;:, -■■■■■• 

fe l« •'■■'v-’V' ' ■' I,;':?-*' :slly ■■,;■■■:' -V. 






'V-..' - -'.r ' ' ' ":. •i^-' . ■' ;^ 










:,;■ ^ .,.:;v. ... r; v^;4 ■ 4' 

- •-■■ *v4: - 1 ' -.: .;V 

^ ^ ^ '■ ' v<j: 


' ■:.;-q - • ,■ ’■. : . ’"V', 'U;:: ■ 

'• v;:'^ ' '4 

_ ■'■'-■■ 4.;vvK'^j^ 

■■ •:.■■■•; 4.' :-i v4-:;.,\:;.. ;-:' .,; ^ 44' ,;!4.^4| 









, ; . ,' ..' ,. ■ ;:v.M 



^4ir S''’^'"' '* • 'V' ' •‘'-'i > * <■ i^‘ . ‘ '■‘-'v' * ' <’ “•^4 - ' ’ ' " '. 



l(l):3-20, 1962 


Journal of Research on the Lepidoptera 

1140 W. Orange Grove Ave,, Arcadia, California, U.S.A. 
Copyright 1962 


VARIATION IN THE SILVERING OF ARGYNNIS 
(SPEYERIA) CALLIPPE IN THE INTERIOR 
MOUNTAIN AREA OF 
SOUTH CENTRAL CALIFORNIA 


OSCAR ELTON SETTE 

23643 Arbor Avenue, Los Altos, California 

In a comprehensive study of the geographic variation of 
Argynnis (Speyeria) callippe Boisduval, based on examination of 
about 2750 specimens from many localities well distributed through 
out the range of the species in California, Hovanitz (1943) recog- 
nized several main divisions of this species complex. Among them 
are the *'South Coast Range Population” extending from the San Fran- 
cisco Bay area southward through the Coast Ranges into Lower Calif- 
ornia and the "Western Sierra Nevada Populations” extending in a 
band along the western Sierra Nevada, at moderate elevations from 
the region west of Lassen Peak southward to about the Greenhorn 
Mountains at the southern end of the Sierra Nevada. He found that 
these two population groups are connected through a "Southern Zone 
of Intergradation” by a "series of steps across the Piute Mountains, 
the Tehachapi Mountains and the Sierra Madre Range.”’ 

He characterizes the populations in the Southern Zone of Inter- 
gradation as follows: 

"From the Santa Monica Mountains ‘ on the coast, where the typical 
callippe of this (the South Coast Range) region lives, it is found that 
in going inland (Charlton Flat, Mint Canyon and ’Ridge Route’) the 
lightness of all colors increases. The butterfly becomes smaller and the 
light colored band and spots on the upper surface of the wings tend to 
become obliterated, leaving a more uniformly colored wing surface such 
as is present in the western Sierra Nevada gradient. However, the yellow- 
brown color is very much lighter than in the latter and the band on 
the under side of the hind wings is still yellow; the spots are still always 
fully silvered. The tendency toward these conditions is the more marked 
the farther from the coast and the farther into the Tehachapi Range 
the populations exist. In the Tehachapi Range, the butterflies are very 
lightly colored and the band on the upper surface of the wings is rare; 
the spots are still silvered. At Havilah, Piute Mountains, the population 
consists of some silvered, some unsilvered and some intermediate spotted 

IThis range is not named on many maps. It lies parallel to, and south of, the middle portion 
of the Cuyama River which is the long north branch of the shorter Santa Mafia River shown 
discharging into the ocean at about lat 35° in Fig. 1. 


3 


4 


SETTE 


J. Res. Lepid, 


individuals (this is the type locality of macaria Edws.); the exact fre- 
quency of these types is not known; but there is a high percentage of 
silvered and unsilvered present ... In the Greenhorn Mountains, the 
segregation into a silvered population on the eastern side of the summit 
and an unsilvered one on the western side is decidedly apparent, though 
mixing occurs toward the south, where the populations unite”. 

The main purpose of this paper is to describe semi-quantitatively 
the variation of silvering within the Southern Zone of Intergradation. 
The silvery decoration of animals is far less common than the pigmen- 
tal coloration. While Hovanitz ( 1941 ) has found general qualitative 
correlations of the pigmental coloration of ^butterflies with environ- 
mental conditions, he finds the significance of silvering in callippe 
"quite incomprehensible" (Hovanitz, 1943, p. 420). Some peculiari- 
ties of the silvering data revealed by the semi -quantitative treatment 
in this paper suggests that studies of its physico-chemical nature and 
genetic control might be rewarding. An hypothesis is advanced as a 
point of departure for such studies. 

THE SAMPLES 

The material for this study consists of complete series of callippe 
that I collected during June 8 to 15, 1957 in company with Fred T. 
Thorne, and one sample from Bouquet Canyon consisting of part of 
a series taken in 1952 by Thorne and kindly given to me. In brok- 
ing the series Thorne exercised no conscious selection for color or 
pattern, though worn and damaged specimens were discarded. 

Parallel or convergent evolution among the Argynnids in some 
regions presents problems of species identification. In the Southern 
Zone of Intergradation this problem arises between calliiype and cor- 
onis Behr. The latter, as the subspecies hennei Gunder, flies with cal- 
lippe and resembles it so closely that confident identification requires 
either great familiarity with both species or known series from the 
region for comparison. L. P. Grey is in possession of both and kindly 
reviewed my field identifications. These had been made in consulta- 
tion with Thorne. At the time neither of us had previous acquain- 
tance with hennei, though we were both familiar with several other 
subspecies of coronis. Two specimens I had labelled callippe proved 
to be undoubted coronis. One labelled callippe came back from Grey 
with the notation, "Could this be coronis? (Don’t ask me!." Another 
labelled coronis bore Grey’s notation "I think it is a callippe!* The 
last two, when spread and compared with the now-available good 
series of hennei, appear to me to be undoubted coronis, though the 
under side wing surface (the only surface exposed to Grey’s examina- 
tion) is remarkably like the callippe from the same locality. These 
four speciments have been excluded from the samples. 

In the list that follows, the collections are grouped by localities. 
Each group contains mainly specimens taken at one location, to 
which have been added a few specimens from nearby locations. Those 


i(j):}-20, 1^62 


VARIATION IN SILVERING 


5 


in the main group, I believe, are members of a single interbreeding 
colony; the others may consist of strays from the same colony in some 
instances, and in others, may be strays from some other colony. All 
localities are in Kern County except Sandberg’s and Bouquet Canyon 
which are in Los Angeles County. Distances from named places are 
given to the nearest mile (I. 6 I km) measured on a straight line on 
large-scale road maps. Directions, also determined from such maps, 
are referenced to true North and stared in Mariners’ abbreviations 
appropriate to the 32 -point compass. Thus southwest is siven as SW, 
southwest by west as SWxW and west southwest as WSW. These are 
11.25° steps and allowing for measuring inaccuracy are correct to 
about the nearest 15°. Elevations were taken by aneroid altimeter 
graduated in 100-foot (30.5 m) units and probably accurate to ± 
200 feet. Botanical names are according to Jepson (1923 - 1925), 
Abrams (1940, 1944 and 1951) and Abrams and Ferris (I960). No- 
menclature of the butterflies follows the list by McDunnough (1938) 
except for the genus {Speyeria), which, at the species and subspecies 
level follows the arrangement of Dos Passos and Grey (1947). 

A. East slope, Greenhorn Mountains. June 12, 1957, 18 males and 
1 female taken 6 miles WSW of Kernville, elevation 4900 ft., along the road 
leading steeply from Isabella Reservoir to Greenhorn Mountain Park passing 
between Tittle and Rattlesnake Creeks, in association of Digger Pine (Pinus 
sabiniana Dough), Oak (Quercus sp), California Fremontia (Premontia cali- 
fornica Torr.), and chaparral broken by grassy areas, with Yerba Santa {Brio- 
dictyon so.) flowers as nectar attractant; included in addition is 1 male taken 
8 miles WxS of Kernville, elevation 5500 ft., Transition Zone, in a small 
roadside opening in Yellow Pine (Pinus ponder osa Dough) and Incense 
Cedar {Libocedrus decurrens Torr.) forest. 

B. West slope, Greenhorn Mountains. June 12, 1957, 58 males and 
6 females taken 27 miles NE of Bakersfield, elevation 3300 ft., near the foot 
of Eugene Grade, in Digger Pine, Blue Oak (Qurcus douglasii Hook & Arn.) 
and grass association, with violets (Viola sp.) abundant; included in addition 
are 3 males and 1 female taken 26 miles NNE of Bakersfield, elevation 3000 
ft., in Blue Oak and grass association, with riparian flora along a small creek. 

C. Havilah. June 11, 1957, 13 males and 1 female taken 2 miles S of 
Havilah, elevation 3100 ft., Digger Pine, mixed oak and chaparral association 
with riparian flora along a small creek, mostly attracted to Yerb Santa flowers; 
in addition are 1 male from 1 mile north of Havilah, elevation 2800 ft., and 
another from 5 miles S of Havilah, elevation 3900 ft. 

D. Walker Basin. June 11, 1957, 30 males and 6 females taken 10 miles 
SxW of Havilah, at the south end of Walker Basin, elevation 3200 ft.. Blue 
Ok and Grass association with sagebrush and violets. Hoarhound (Marrubium 
vulgare L.) attracted a few individuals, but most were taken in the characteristic 
slow, fluttering flight displayed when on breeding grounds. 

E. Tehachapi. June 14, 1957, 12 males and 7 females taken 8 miles W 
of Tehachapi (the town), elevation 4700 ft, at the eastern end of Bear Valley 
in Blue Oak and grass association, with patches of sagebrush (Artemisia tri- 
dentata Nutt.), mostly acctracted to Wallflower (Erysimum sp.) flowers; in 
addition are 3 males from 7 miles W of Tehachapi, elevation 4600 ft., and 1 
female from 3 miles SW of Tehachapi, elevation 4600 ft. 


6 


SETTE 


/. Res. tepid. 


I2 0‘ 


119 * 


118 ® 



36 ' 


35 * 


3 4 * 


FIGURE 1. Location of populations sampled (letters "A to H”) super- 
imposed on the distribution of Argynnis (Speyeria) callippe (shaded area) 
adapted from a portion of Hovanitz’ map (1943, Fig. 1) showing the distribu- 
tion of the "silvered spots” character by shading lines slanted upward to the 
left and the "unsilvered spots” character by shading lines slanted upward to 
the right. The Kern River (upper right) separates the Greenhorn Mountains 
to the north from the Piute Mountains to the south as it flows to a sink in the 
lower San Joaquin Valley (blank area at upper middle). At middle left is the 
Santa Maria River with the Cuyama River as its long north fork. At the lower 
right is the Santa Clara River. The crests of the higher mountains and ridges 
of the transverse series of ranges lie north of the Santa Clara River Drainage and 
south of the Cuyama River. 


F. Lebec, June 8, 1957, 24 males and 8 females; June 9, 1957, 26 males and 
4 females; and June 15, 1957, 11 males and 15 males; total, 61 males and 
27 females taken 1 mile S of Lebec, elevation 3600 ft., in oak and grass 
association with adjacent chaparral including sagebrush {A. tridentata) , Buck- 
wheat Brush {Eriogonum^ fasciculatum Benth.) and Yerba Santa; some were 
at flowers of the last, others in breeding-ground flight. 


(r;.-j- 20 , 1^62 


VARIATION IN SILVERING 


7 


G. Sandberg^S (on old Ridge Route). June 9, 1957, 12 males and 3 females; 
June 15, 1957 , 1 male and 2 females; total, 13 males and 5 females taken 9 
miles ESE of Gorman in Digger Pine and sagebrush (not A. tridentata) associa- 
tion, with violets. Breeding ground behavior was predominant. 

H. Bouquet Canyon. June 11 and 12, 1951, 18 males collected by Fred 
T. Thorne at "upper Bouquet Canyon, 4 miles down from Pine Creek — Lake 
Hughes road” (estimated 16 miles W of Palmdale, estimated elevation 
3400 ft.). 


HABITAT AND MESOSCALE DISTRIBUTION 

According to these records callippe in this region occupies ele- 
vations from 2800_ to 5500 feet, but occurs in good concentrations only 
between 3100 and 4900 feet, generally, the upper portion of the Up- 
per Sonoran Life Zone. Its habitat is marked characteristically by scat- 
tered Blue Oak, Digger Pine, and sparse chaparral with sagebrush {A. 
tridentata) often pesent. Usually there are broad areas or small patches 
of grass present. Practically all of the localities are in grazing land, 
but at the time of our trip it was being very lightly grazed. 

Violets were noted at only three of the locations. Where they were 
seen, the plants were abundant, mature, with fully-developed seed cap- 
sules and appeared about to dry up. Where violets were not seen, 
either they had dried up already, or the collecting was of members of 
the colony that had been drawn by nectar sources somewhat away 
from the site of the larval food plant. 

During the trip we collected at 31 locations within the geographic 
range and elevation limits of this butterfly. Of these, seven obviously 
were in, or close to, callip'be colonies; five locations yielding only one 
or several specimens, evidently were not as close; and the remaining 
19 locations where we took none probably were distant from colonies, 
though this may not have been true for a few places where we sighted 
an Argynnid or two v/hich could have been either this species or coro- 
nis. These indications favor a population model consisting of well- 
separated, compact colonies from which individuals do not stray often 
or far. This pattern is more accentuated in this region than in most 
others within my collecting experience. 

Butterflies of 30 other species were taken at one or another of the 
seven locations where callippe were caught in good numbers. Plebeius 
acmon West, and Hew. and Hesperia lindseyi Holland were the most 
ubiquitous, occuring at most locations from the Greenhorn Mountains 
to Sandbergs. The former were taken in small numbers while the latr 
ter, in some locations were very abundant. In addition to these, the 
most common and prevalent in the Greenhorns and the Piutes were 
Euphydryas chalcedona Doubleday and Hewitt, Melitaea palla Boisdu- 
val, and Strymon saepl^m^ Boisduval; while through the Tehachapi to 
Sandberg’s they were Minois silvestris Edwards and Argynnis coronis 
Behr. If one of the specimens whose identification was questioned by 


8 


SETTE 


/. Res. Lepid. 


Grey is, as I believe, coronis, this species extended north at least to 
Havilah. Inasmuch as all coronis taken on this trip were fresh males, 
and in other regions within my collecting experience coronis flies some- 
what later than callippe, it is likely that a week or two later in the 
season coronis would have been found to be even more consistently 
a companion of callippe. 

RELATIVE ABUNDANCE 

The relative abundance in butterfly populations may be signifi- 
cant in studies of variation. From records extending through many 
years, Ford (1945) found that a rapid increase in abundance of the 
colonial butterflv Euphydryas aurina Rott was accompanied by "an ex- 
traordinary outburst of variation". When the increase ceased, the 
colony settled down to a comparatively uniform type, different from 
the one prevailing prior to the increase. Scientists differ as to the in- 
terpretation and significance of this phenomenon as a mechanism in 
evolution. Whatever the interpretation, if the generality of this phen- 
omenon is to be ascertained, records must be accumulated through the 
years for a number of butterfly populations. 

To this end, there are placed on record in Table 1, estimates of 
relative abundance of callippe in each of the seven colonies sampled 
in 1957. This estimate is in terms of catching rate and assumes that 
the number caught in unit time is proportional to the abundance of 
the population in the area of collecting. It is homologous in concept 
to the measure "catch per unit of effort” almost universally and suc- 
cessfully (sometimes "calibrated” to a absolute abundance by tag-and- 
recapture experiments) used as a basic statistic in studying the dy- 
namics of fish population fluctuations (Ricker 1940). For callippe 
I have computed the catching rate, R, according to the formula: 

R ^ Nc . 

H - k Nt 

where Nc is the number of callippe caught, H is the time, in hours, 
in the collecting period, Nt is the total number of specimens of all 
species caught (including callippe) during the collecting period, and 
k is a constant whose value depends on the methods and dexterity of 
the collector in caring for a specimen once it is in the net. By timing 
myself in the_caring, individually, for each of 150 specimens of a 
number of species from all families except Megathymidae under typ- 
ical field conditions, I found my k value to be 0.0155 hours (37.8 
seconds ) . 

In repeated samplings of several species I have fo*und R to be sur- 
prisingly stable. For callippe, the three samplings at Lebec are an ex- 
ample of this. Although stable, R as computed by this formula should 


i(i):3’2o, I $62 


VARIATION IN SILVERING 


9 


be regarded only as a first-order approximation, or index, of true rel- 
ative abundance, because it is affected by systematic error and several 
sources of variability, not all of them random. A more sophisticated 
iterative treatment theoretically should improve R, but it involves as- 
sumptions of uncertain merit awaiting test. 

TABLE 1. Relative abundance in terms of catching rate, R, in number per 
males and by the per cent of the males that were fresh. 


Catch- 
ing Per Per cent 

Locality, date (1957) and elevation rate cent of males 

(R) males fresh 


A. E. slope, Greenhorns, el. 4900 ft., June 12 ....... * 39 84 86 

B. W. slope, Greenhorns, el. 330 ft., June 12 ........ **227 91 52 

C. Havilah, el. 3100ft., June 11 16 93 15 

D. Walker Basin, el. 3200 ft., June 11 * 44 83 87 

E. Tehachapi, el. 4600 - 4700 ft., June 14 33 67 79 

F. Lebec, el. 3600 ft., June 8 24 75 84 

F. Lebec, el. 3600 ft., June 9 24 87 69 

F. Lebec, el. 3600 ft., June 15 30 42 45 

G. Sandberg’s, el, 4000 ft., June 9 17 80 75 


In the meantime, of most concern is that R increasingly underes- 
timates abundance as the latter increases, presenting simultaneous 
catching opportunities more often and usually avail can be taken of 
only one of them at a time. Recognizing this, the values in Table 1 
that probably were moderately underestimated owing to this factor 
are marked with an asterisk, and the one value that was grossly under- 
estimated is marked with a double asterisk. At the other end of the 
scale, when only one or two specimens are caught during a collecting 
period, R has large error owing to random variability of incidences 
of encounters. Avoiding this. Table 1 gives catching rates computed 
for only the first-listed location under each locality, except for Tehach- 
api where the times and catches of the first two locations were pooled 
for computing the catching rate. 

Another error source is that as kNt approaches H, a small error 
in k produces a large error in R. This approach was not close enough 
to be critical for the values given in Table 1, though the value for the 
west slope of the Greenhorn Mountains may have been moderately 
affected. 

Every collector can think of a number of other obvious sources of 
error or variability such as weather conditions, unusual concentrations 
at attractants, etc.. These obvious ones can be avoided by comparing 
only sets of samplings taken under reasonably similar conditions, as 
is true for the set in Table 1 except as later noted. 

Of course the catching rate reflects the relative abundance only at 
the stage of flight at which the collecting was done, and may yield 
values departing substantially from the inherent abundance. An esti- 


10 


SETTE 


/. Re$. tepid. 


mate of the stage of the flight is afforded by the relative numbers of 
fresh and worn individuals. In Table 1, recorded as *'fresh” were in- 
dividuals without readily perceptible random loss of scales or fringe. 
Damage reasonably attributable to accidental causes, such as notches, 
tears, breaks and rubbed streaks or patches was not considered. The 
data are given for males only, there not being enough females to yield 
reliable percentages. The two sexes were not combined for this sta- 
tistic because this would introduce extraneous variability owing to the 
tendency of females to emerge later than males. As evidence of this, 
only four of the 53 females listed in Table 2 were worn. More direct 
evidence is afforded by the Lebec samples. Those of June 8 and 9 were 
mostly males and mostly fresh. About a week later more than half of 
the males were worn and the females, all fresh, outnumbered the males. 
The sample from Havilah is anomalous in respect of percentage worn 
in relation to sex ratio. Although most of the males were worn, only 
one female was taken. Probably the greater ease of collecting males at 
the Yerba Santa flowers diverted us from the females which probably 
were widely scattered for ovipositing in the adjacent area which ap- 
peared to be the likely habitat for violets, though the plants were not 
observed, doubtless having dried up before our visit. At Sandberg’s 
the catching rate was substantially depressed bv a strong breeze sweep- 
ing the exposed slope and wafting many a disturbed butterfly out of 
stalking or pursuit range. 

Appraising the catching rates in the light of these qualifying fac- 
tors, the presence of some worn males at all localities indicates that the 
callippe flight was well developed throughout the region. But the high 
percentage of fresh males and the scarcity of females suggests that the 
flight had not yet reached its peak at most localities except Lebec where 
it probably was near peak stage on June 15 and Havilah, where the 
male flight had apparently passed its peak. Allowing for the differ- 
ences in stage of flight and the collecting difficulties at Sandberg’s, it 
appears that relative abundance was capable of attaining a height-of- 
flight index of 30 to 40, probably closer to the latter, at all localities 
except the location on the west slope of the Greenhorn Mountains 
where abundance was at least an order of magnitude higher, and at 
Walker Basin where it probably was substantially higher than 40 per 
hour, though well below that on the west slope of the Greenhorn Moun- 
tains. Although we did not trace out the geographic extent of a colony 
in any of the localities, the impression gained while collecting, was 
that these two colonies covered more extensive areas than the rest. It 
may be concluded, first, that the samples were drawn from near the 
middle of the flight period and hence probably represent nearlv the 
modal characteristics of the population in each colony; and second, the 
colonies varied quite widely in population size, perhaps through more 
than one order of magnitude. Further results may emerge if compar- 
isons with changed levels of abundance become possible in the future. 


1^62 


VARIATION IN SILVERING 


11 


VARIATION IN SILVER SCALING 

The silver scaling in callippe^ as in Argynnids generally, is confiined 
to certain pattern elements on the ventral wing surface. For the most 
part these are well defined spots in the apical and subapical area of 
the fore wing and all well defined spots in all areas of the hind wing. 
In addition, on heavily silvered individuals, silver scales may form ill 
defined streaks along the inner margin and in some of the interspaces 
between spots in the basal and discal areas of the hind wing. The pres- 
ent study is confined to the well defined spots of the hind wing. When 
unsilvered these spots are a light buff color, usually sharply bordered 
or outlined by brown scales. Some or all of the spots may be partly sil- 
vered. Then the silver scales and buff scales stand out in sharp con- 
trast to each other when specimens are held in the best relation to the 
light source and the eye to bring out the specular quality of the silver 
scales. 

A fully quantitative measure of the degree of silvering would be 
the ratio of silvered scales to all scales in the area potentially subject 
to silvering. To avoid spending a prohibitive amount of time count- 
ing scales, I have employed a subjective system of scoring based on 
general appearance. It would have been desirable, too, to utilize ex- 
clusively the pristine fresh individuals, but to avoid reducing the num- 
ber below levels needed for tests of significance, I have scored the 
slightly and moderately worn individuals along with the fresh ones, ex- 
cluding only those whose loss of scales caused serious doubt as to their 
correct score. The penultimate column of Table 2 shows the number 
excluded for this reason. 

Because the amount' of silvering sometimes differed appreciably in 
the basal and discal area, which will be called "disc” for brevity, from 
that in the submarginal area, called "margin”, the two portions of the 
wing were scored separately. Four grades of silvering were recognized: 
Grade 0 for complete lack of silver scales: Grade 1, when there were 
only flecks of silver well separated by buff scales; Grade 2, when some 
of the spots were unsilvered, others were partly silvered and still others 
were fully silvered, or any combination of two of these conditions; and 
Grade 3, when all spots were so fully silvered that there were no readily 
perceptible buff scales. The Disc Grade and the Margin Grade were 
treated as half scores and added together to arrive at the individual 
Score. Thus there are seven possible scores: 0 to 6, inclusive. 

Of the 238 individuals scored, 50 had half scores that differed from 
each other by one grade point. None differed by more than one. Of 
the 50 with different half scores, the Disc Grade was higher than the 
Margin Grade for 40 individuals and lower for 10. Curiously, for the 
two Greenhorn Mountain samples and the' Havilah sample, pooled, the 
ratio was 33:4, while for the Walker Basin, Tehachapl and Lebec sam- 
ples, also pooled, it was 7:6. The difference between the ratios 33:4 


12 


SETTE 


/, Re$. tepid. 


and 7:6 is statistically significant (p=0 02) desoite the small numbers 
in the latter. It is possible that the tendency for the marginal row of 
spots to be less silvered than the disc spots is linked to the general lack 
of silvering in the three northernmost localities. 

TABLE 2. Frequency distribution of silvering scores 

Sex and sample Score 

0 1 ' 2 3 4 5 6 Mean Scored Total 


Males : 

A. E. slope. Greenhorns 5 3 1 1—2 1 3.16 — - 19 

B. W. slope, Greenhorns 17 8 7 64 8 7 2.42 7 64 

C. Havilah 2 1 — 2 — 2 4 3.73 4 15 

D. Walker Basin 1 2 1 — 1 2 19 5.08 4 30 

E. Tehachapi 1 — — - i— _ — 13 5.40 — 15 

F. Lebec 1 1 1 1 1 1 39 5.54 15 60 

G. Sandberg’s — — — — — . — 10 6.00 3 13 


Total 27 15 10 11 6 15 99 4.16 33 216 


Females : 

A. E. slope, Greenhorns — 1- — ■ — — — -2.00 — ■ 1 

B. W. slope. Greenhorns — — 2 3 - — — — 2.60 5 

C. Havilah l l 

D. Walker Basin _ „ _ 11 1 3 5.00 — 6 

E. Tehachapi _ _ l 1 6 5.01 — 8 

F. Lebec 1 _ _ _ 25 5.77 1 27 

G. Sandberg’s - — — — . — — — 3 6.00 — 5 

Total 1 — 4 5 1 i 39 5.12 2 53 


The mean score for males is 4.16 and for females is 5.12 (Table 2), 
suggesting that females tend toward more silvering than males. But 
four-fifths of the females were taken at Walker Basin, Tehachapi and 
Lebec where silvering in both sexes is nearly complete. If we pool 
these three sampls and cHss together the individuals with scores 0 to 
4 inclusive, and similarly those with scores 5 and 6, in order to have 
high enough numbers for statistical test, we find no significant dif- 
ference between the sexes (P = 0.9). Accordingly, further analysis 
deals with both sexes combined, as shown graphicallv in Fig. 2. 

The arrangement of samples in Fig. 2 is in ascending order of their 
means. This also arranges them from north to south except for the 
reversal of samples A and B. Where samples are near the same par- 
allel of Latitude and have appreciable meridianal displacement, as 
among A, B and C as one group and F, G, and H as another, their 
means also ascend from west to east. 

The slopes of the lines connecting the means in Fig. 2 should not 
be taken as representing gradients of silvering in the sense of reflecting 
unit increase of silvering per unit distance. The panels are not spaced 
in proportion to distance, but merelv to accommodate the height of the 
bars. The means, themselves, are defective because the scores do not 
represent even gradations in amount of silvering. Further, because of 


t§€a 


VARIATION IN SILVERING 


13 


wesr SLO PE. GREEN - 
HORN MTS. 35 * 37 ' 
EAST SLOPE, GREEN - 
HORN MTS. 35 ® 43 ' 

HAVILAH 35 ® 31 ' 


WALKER BASIN 

3 5 * 23 ' 


TEHACHAP! 35 ® 08 ' 


LEBEC 34 * 50 * 


SA NDBERG'S 
34 ® 45 ' 


BOUQ UET CANYON 
34 ® 35 ' 



FIGURE 2. The degree of silvering of the several samples, shown as histo- 
grams of the number of individuals according to grade (see text); and the 
mean grades, shown as small circles connected with heavy straight lines. 


NUMBER OF INDIVIDUALS 


14 


SETTE 


J. Res. Lepid. 


the prevalence of low numbers at the middle scores, the distributions 
depart widely from the bell-shaped normal probability curve and the 
par? metric test commonly applied to means would not yield reliable 
probabilities even if the means were not otherwise defective. The 
means were computed and connecting lines were supplied in the graph 
only to give a general perspective of the amount and direction of dif- 
ferences between localities. 

For a more careful assessment, the significance of the differences 
between samples, as to the numbers of individuals falling into the sev- 
eral silvering classes, can be made by the chi -square test which re- 
quires neither a normal distribution nor uniform class size. It does 
require a pooling of the original score classes to provide a minimum 
of five as the expected number on a class. To meet this requirement, 
the data have been grouped into two classes: (1) individuals with 
Scores 0 - 4, which can be described as "unsilvered” and partly silvered”; 
and ( 2 ) those with Scores 5 and 6, which can be described as "silvered 
or nearly so”. With this grouping the approximate probability, P. of 
the difference between any two samples in respect of the ratio of "un- 
silvered and partly silvered” to "silvered or nearly so” occurring by 
chance can be found by entering the table of chi-square distribution 
on one degree of freedom with the value of chi scfuare computed from 
the 2x2 contingency table according to the method given by Fisher 
(1948, p. 85). For greater precision the Yates correction for contin- 
uity was incorporated and P was found by graphical interpolation be- 
tween tabular values. The results are given in Table 3 as a matrix in 
which the chi-square values betyeen all combinations of pairs of sam- 
ples are shown in the upper right portion and the corresponding values 
of P in the lower left portion. 


TABLE 3. Chi-square values (upper right) and probability values (lower left) 
between pairs of samples. 


Locality 




Sample 





and sample 


B 

A 

C 

D 

E 

F 

G 

W. Greenhorns .. 

B 


2.37 

2.97 

>10 

>10 

>10 

>10 

E. Greenhorns .. 

A 

0.16 


0.02 

4.63 

1.10 

>10 

7.14 

Havilah 

C 

0.08 

0.10 


1.24 

1.74 

0.83 

1.78 

Walker Basin 

D 

< .001 

.03 

.19 


.01 

.84 

2.32 

Tehachapi 

E 

<.001 

.02 

.18 

.10 


.64 

1.36 

Lebec 

F 

<.001 

<.001 

.34 

.34 

.40 


.37 

Sandberg’s — 

G 

<.001 

.008 

.02 

.14 

.19 

.10 



The three northern samples. A, B, and C form a group within which 
there is no statistically significant difference at the five per cent prob- 
ability level (P = 0.05), and the four southern samples, D to G, in- 
clusive, form another such group. But these groups are not discrete 
from each other. Sample C from Havilah differs no more significantly 
from D, E, and F than from A and B, indeed, somewhat less. The sum 
of chi squares among A, B and C is 5.36; df are 3; and P is 0.14. The 


i(i):3-20, 1962 


VARIATION IN SILVERING 


15 


comparable values among C, D, E, and F are: Chi-square, 5.34; df, 6; 
P, 0.48. The indeterminate position of Havilah relative to the two 
groups, most probably is due to the small number in the sample from 
there. With such a small sample it would require a very strong con- 
trast in silvering ratio to prove statistical significance. Until more ma- 
terial is available it remains uncertain whether the population at Havi- 
lah resembles more closely those to the North or those to the South in 
respect of silvering, or whether it is truly intermediate as suggsted by 
Fig. 2. 

Likewise, the sample from the east slope of the Greenhorn Moun- 
tains is small and does not test significantly different from that of the 
west slope. But for the Greenhorn Mountains there are two sources of 
additional data. Thorne and I scored both his catch and mine while 
in the Greenhorns and this record includes about twice as many indivi- 
duals as given in Table 2. We used three categories: ' unsilvered”, 
*'part silvered”, and “silvered”, corresponding somewhat inexactly to 
the Scores 0 - 1, 2 - 4 and 5 - 6, respectively, of Table 2. Hovanitz 
(1934) reported the number of individuals in a sample from the east 
side of the Greenhorn Mountains, elevation 5500 feet, and from the 
west side at Cedar Creek, elevation 5000 feet, according to three cate- 
gories: “not silvered or very slightly so”, “intermediate” and “well - 
or fairly-well silvered”. Assuming these categories correspond approx- 
imately to those of Thorne-Sette, the data may be pooled as follows: 
East side: 



Unsilvered 

Intermediate 

Silvered 

Total 

Thorne - Sette . 

......6 

.15 

17 

38 

Hovanitz 

....0 ...... 

..2 

14 

16 

Total 

West side: 

...........6 

17 ......... 

31 

...54 

Thorne - Sette ... 

72 

...24 

32 

128 

Hovanitz 

....10 

8 

4 

22 

Total .............. 

.........82 ...... 

..........32 

36 

150 


With more data it is possible to use a 2 x 3 contingency table for 
a more discriminating test. This yields a probability far less than one 
in a thousand that the west side and east side samples could have been 
drawn from a population containing the same proportions of unsil- 
vered, partly, silvered, and silvered members. This does not prove 
that there is no mixing between the populations of the west and east 
slopes; it only indicates the extreme unlikelihood of enough inter- 
change to form a thorough mixture. But when it is considered, in 
addition, that the ridge of the mountain is clothed with transition 
forest within which no colonies of callippe were found, it seems clear 
that the interchange, if any, must be slight and probably by way of 
mixing with the more silvered populations to the South at different 
rates on the two sides of the mountain range as suggested by Hovan- 
itz (1943, p. 411). 


16 


SETTE 


/. Res. tepid. 


Turning southward, past Havilah, there is at Walker Basin a small 
proportion of unsilvered and partly silvered individuals in the pop- 
ulation. The low proportion continues with only slight further dim- 
inution in going from Walker Basin in the Piute Mountains, through 
the Tehachapi Mountains to Lebec. This uniformity suggests a rela- 
tively brisk interchange between colonies in this stretch of mountains, 
probably coupled with strong environmental selection against unsil- 
vered individuals. 

Then at Sandberg’s, only a short distance beyond Lebec, all indiv- 
iduals are silvered, This is true also for the specimens from Bouquet 
Canyon (not included in the scoring table because they were only a 
part of a series). But my sample from Sandberg’s is too small to 
assuredly include unsilvered or partly silvered individuals if they con- 
stitute only a small percentage of the population. There is historical 
evidence, in Gunder’s (1930) list of butterflies of Los Angeles Coun- 
ty, of the occurance of unsilvered callippe at several localities in the 
transverse ranges south and southeast of Lebec. Under ”Araynnis ma- 
caria lamina Wri. THE UNSILVERED MACARIA FRITILLARY, a 
transition form.” Gunder recorded one male collected by Comstock 
"June 10, 1922, Ridge Route;” one male collected by Friday "June 19, 
1929, Pine Canyon;” and two males and one female collected by Gun- 
der "June 25, 1921, Bouquet Canyon.” "Ridge Route” probably is 
identical to, or at least near, the locality I give as "Sandberg’s”; "Bouquet 
Canyon” may be identical to, or near the locality of the Thorne col- 
lection here listed under the same place name; Pine Valley lies about half- 
way between the two. The last is definitely on the desert side of the 
transverse ranges, while the other two localities are near the crest divid- 
ing the drainage to the desert from the drainage to the Pacific. The 
Thorne collection is definitely from the Pacific drainage side. It is proba- 
bly that this crest marks the southern limit of the unsilvered or partly 
silvered form. 

Reviewing, there is a mixture of unsilvered, partly silvered and 
fully silvered callippe extending through the Southern Zone of Inter- 
gradation, from the Greenhorn Mountains in the North, southward 
through the Piute and Tehachapi ranges, to the crest of the series of 
ranges extending transversely across California at about latitude 34° 
40’ N. Most of the change from the predominantly unsilvered con- 
dition in the north to the predominantly silvered condition in the 
South, take place in about the northernmost one-fourth of the Zone. 
Through the remaining three-fourths, beginning at about the mid- 
length of the Piute Mountains, where a high degree of silvering al- 
ready has been reached, the increase in silvering is very gradual. Al- 
though it is still perceptibly incomplete at Lebec near the Southern 
end of the Tehachapi, unsilvered and partly silvered become so rare 
beyond there that one should not expect to encounter evidence of the 
unsilvered condition without drawing large samples from the popu- 


(i):}-20, ip 62 


VARIATION IN SILVERING 


17 


lation. It is reasonable to conclude that this is the southern limit of 
the tendency toward lack of silvering in the ventral hind-wing spots 
of callippe. To express this finding in Fig. 1, the area shaded with lines 
slanted upward to the right should be extended to include localities 
E, F, G and doubtfully H. When so extended, the southern limit of 
lack of silvering would more nearly agree with the southern limits of 
"uniformly colored red-brown upper surface” and "lack of brown pig- 
ment between sdo<-s on the underside of the hind wings” as mapped 
by Hovanitz (1943, Figs. 2 and 4). 

DISCUSSION 

Although this description adds some details to the distributional 
pattern of silvering in callippe, it raises more questions than it an- 
swers. Some are concerned with geographic limits. How much far- 
ther north does silvering extend? Does it taper off gradually from 
the western Greenhorn Mountains northward as does the opposite 
condition from Walker Basin southward, or does it end abruptly a 
short distance north of the sampled Greenhorn Mountain localities? 
Does the more silvered population of the eastern slope of the Green- 
horn Mountains end as in a cul-de-sac co-terminal with the limits of 
the Kern River drainage, or does it extend northward as a tongue 
along the higher Sierra Nevada toward silvered populations such as 
those near Downieville and Gold Lake? 

More perplexing questions concern the processes governing silver- 
ing. The distribution of individuals among the seven score classes re- 
veal a paucity of intermediates in the middle of the range (histo- 
grams A and C in Fig. 2 ) . Could such a frequency distribution arise 
from simple Mendelian inheritance, or is a more complex mechanism 
required? 

The existance of intermediate deo:rees of silvering suggests sim- 
ilarity with the example of simple Mendelian inheritance described 
by Ford (1945, p. 173) for the Currant Moth, Abraxas grossulariata 
L., which, in its commonest form, has wings with a white bick.Erround. 
In its less comomn form, lutea Cockerell, the white is replaced by a 
deep yellow. When these two forms are interbred the heterozygotes 
have a pale yellow background. Mating an individual homozygous for 
white with one homozygous for deep vellov/, produces offspring inter- 
mediate between the two parents. There is no dominance. Taking 
this as a model for silvering in callippe, and using Ford’s system for 
notation we may use the symbols and for represen- 

ting the unsilvered ( Score 0 ) , the partlv silvered ( Scores 1 - 5 ) , and 
the completely silvered (Score 6) phenotypes, respectively. For such 
a model, in a population containing equal numbers of the two homo- 
zygotes (C^C^ and OC^), we should find : C^C^: 

1:2:1. None of our samples have equal numbers of homozygotes, 
but if we pool samples A, B and the ratio is 24 : 51 : 18. This 


18 


SETTE 


/. Res. Lepid. 


differs from the ratio 1 : 2 : 1 by no more than one would expect 
from random sampling variability in about half of the samples drawn 
(P =z 0.44). We may conclude that there is no evidence against the 
hypothesis that silvering is controlled by a single pair of alleles of the 
phenotypes scored 1 to 5 may be lumped as the heterozygous com- 
ponent of the population. 

However, in the Currant Moth, equal doses for white and for 
deep yellow produced a color about half way between the two homo- 
zygotes. In callippe there are very few individuals about half silvered. 
Individuals scored 3 and 4 comprise less than one-third of the cate- 
gory with scores 1 to 5. Perhaps some more complex genetic system 
may account for this peculiar distribution within the partly silvered 
segment of the population. But I am attracted by the idea that some 
chemico - physical process, such as crystallization may be involved. 
These tend to be triggered by very slight differences in conditions, 
and once triggered, go to completion. 

In this connection it is also interesting that in areas of potential 
silvering, the single scales, as viewed with a hand lense, appear either 
totally buff or totally silver, never in between. 

In pursuing this idea, I have found only a little information on 
the silvery decorations of the Lepidoptera in the literature that bear 
on its chemical nature. Mayer (1897) makes direct reference to its 
nature in the Argynnids, saying ”... Dimmock ('83) has shown that 
the silvery white and milk-white colorations are due to optical effects 
produced by reflected light. In the silvery white scales, however, such 
as the under surface of the hind wings of Argynnis, there must be 
a reflecting surface toward the observor, for both silvery and milk- 
white colors appear simple milk-white by reflected light.” According 
to Fox (1953, p. 289) uric acid, derived from chrysalid metabolism, 
deposited in the wings of the adult butterfly; guanine and uric acid 
contribute opaque whiteness and glistening silvery aspects. Ford 
(1945), although discussing extensively the physical and chemical 
nature of butterfly coloration, is silent on the nature 'of silvering. Taylor, 
(1925) points out that guanine (also spelled guanin) deposited in 
crystalline form imparts the silvery appearance prevalently displayed 
by pelagic fishes, while bottom fishes with prevalently white under- 
surfaces have their "subdermal tissues heavily charged with amorphous 
guanin, which is chalky white.” He describes guanine crystals, after 
they have been processed into pearl essence for making artificial pearls, 
as ". . . very thin light blades, floating in a liquid . . . [which] show 
their maximum luster when they are oriented parallel to each other 
. . . [when] pointing promiscuously in all directions, the effect will be 
a metallic or dull pearly luster.” 

While one would prefer evidence derived directly by analysis of 
the substance as it occurs in callippe, it is not far fetched to suppose 
that this substance is guanine, sometimes deposited as crystals disposed 


t(i):3-20, i$6a 


VARIATION IN SILVERING 


19 


with appropriate orientation to give the specular effect that we call 
silvery, and sometimes in amorphous form adding whiteness to the 
pigmental brown to produce the light buff of the unsilvered spots. 
On this supposition it is necessary that the "silvering gene” control 
only the conditions within the pupal scale-sac fluid so as to precipitate 
guanine amorphously for buff scales, or alternatively, as crystals for 
the silver scales. Presumably, in homozygotes the balance of influences 
is well to one side of the critical point so that the deposite in all 
scales is always amorphous, or always crystalline. In the heterozygotes, 
too, it may take place on the "all or none” basis, but by single scales. 
Under the equal and opposite influence of the two alleles the equilib- 
rium may be very unstable and readily tipped to one side or the other 
of the critical point. This could be mediated by very slight differences 
in environmental conditions of the particular microclimate of a pupal 
individual at the precise time of color deposition in the scale sacs. 
This would tend to affect nearly all of the scales of an individual alike, 
swinging most of them toward one side of the critical point more often 
than nearly equal numbers to each side. Small differences between 
members of the colony as to pupation site, date and time of day of 
deposition of salts in the wing sacs, or even the rate of drying after 
eclosion, might provide the variety of conditions necessary to produce 
the observed peculiar frequency distribution of the supposed hetero- 
zygous phenotypes. 

This set of ideas is not advanced as an explanation, but as an 
hypothesis inviting test by those having interest and competence in 
the fields of experimental breeding or biochemistry, or both. 

It is a pleasure to acknowledge the stimulus to undertake this 
analysis and the material help received from William Hovanitz; the 
generosity of Fred Thorne in giving specimens and sharing his uncom- 
mon collecting serendipity; and the kindness of Paul Grey in review- 
ing my identification of many hundreds of specimens of this and other 
Argynnids. 

LITERATURE CITED 

ABRAMS, LEROY. 1940-1951. Illustrated Flora of the Pacific States, vols. 
1-3, 4 vols. Stanford. 

ABRAMS, LEROY and R. S. FERRIS. I960. Illustrated Flora of the Pacific 
States, vol. 4, Stanford. 

DOS PASSOS, C. F. AND L. P. GREY. 1947. Systematic Catalogue of 
Speyeria with designation of types and fixation of type localities. Amer. 
Mus. Novitates, no. 1370: 1-30. 

JEPSON, W. L. 1923-25. A Manual of the Flowering Plants of California. 

Berkeley. . 

FISHER, R. a. 1948. Statistical Methods for Research Workers. Hafner, 
N. Y. 10th Ed. 

GUNDER, J. D. 1930. Butterflies of Los Angeles County, California. 

Bull. Sou. Calif. Acad. Sci. 29 ( 2 ) ; 39-95. 

HOVANITZ, WILLIAM. 1941. Parallel ecogenotypical color variation in 
butterflies. Ecology 22: 259-284. 


20 


SETTE 


J. Res, Lepid. 


1943. Geographical variation and racial structure of Argynnis callippe 
in California. Amer. Nat. 77:400-425. 

MAYER, ALFRED G. 1897. On the color and color-patterns of moths and 
butterflies. Bull. Mus. Comp. Uool. Cambridge 30 (4) : 169-256. 

RICKER, W. E. 1940. Relation of "catch per unit effort” to abundance 
and rate of exploitation. /. Fishery Res. Board Canada, Toronto. 

TAYLOR, HARDEN F. 1925. Pearl essence: its history, chemistry, and 
technology. Appendix to: Rept. U.S. Commissioner Fisheries for 1923. 
Bur. Fish. Doc. 989. Washington, D.C. 


1 ( 1 ) : 21 . 42 , 1962 


Journal of Research on the Lepidoptera 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
Copyright 1962 


THE EFFECT OF VARIOUS FOOD PLANTS ON 
SURVIVAL AND GROWTH RATE OF PIERIS^ 


WILLIAM HOVANITZ AND VINCENT C. S. CHANG 

California Arboretum Foundation, Inc., Arcadia^, Los Angeles State College, 

Los Angeles and University of California, Riverside 


The close relationship between most phytophagous insects 
and their host plants is a general phenomenon known by most 
entomologists. This relationship is so outstanding that field naturalists 
can often locate colonies of any particular insect most readily by first 
searching out the host plants, as they often are larger and more obvious 
to the eye than the insects themselves. 

What is not so well known is ( 1 ) the extent to which it is possible 
for any particular species or race to survive on various food plants, 
and (2) the causes of the attraction to, and survival of, any race of 
insect on a particular plant. 

It is the purpose of this paper to indicate something of the range 
of ability to survive of two insects, Pieris rapae and Pieris protodice, 
(the "cabbage or mustard "butterflies) on various cruciferous plants. 
In other papers, it is intended to go into the various aspects of the 
development of food plant preferences in insects, including its causes, 
evolution, and relationship to origin of phylogenetic groupings. 

THE MATERIAL AND METHOD 

The experimental work to be reported on here and in subsequent 
papers has been carried out in Arcadia, Southern California. All species 
of the genus Pieris (Lepidoptera: Pieridae) found in southern Cali- 
fornia have been utili 2 ed for the study. The geographical ranges and 
specific food plants of the species existing in southern California are 
being described elsewhere (Hovanitz, 1962) and the general distribu- 
tion of the species of North America in another paper (Hovanitz, 
1962). Five species of Pieris exist in this area, P. rapae (a European 
immigrant), P. protodice, P. beckeri, P. sisymbrii and P. napi. Each 
of these occupies a specific habitat and is restricted to a different food 
plant.^ 

1 Aided by a grant from the National Science Foundation, Washington, D.C. 

2The authors wish to express their great appreciation to Dr. W. S. Stewart, Director, Los Angeles 
State and County Arboretum for great cooperation in making this work possible. 

3Some taxonomists consider Pieris occidentalis to be a distinct species separate from Pieris protodice, 
rather than a geographical race, ecological race or local phenotypic form. If this were the 
case, there would be six species in southern California. 


21 


22 


HOVANITZ AND CHANG 


/. Re$. Lepid. 


Pieris rapae is the easiest species to breed under greenhouse condi- 
tions and has been used for most experiments. It aslo has a wider 
range of satisfactory food plants than the others. As a European 
immigrant, it remains restricted largely to plants of European origin 
and has never adapted itself under natural conditions to any American 
native. Contrariwise, Pieris protodice, a native species, has never 
adapted itself to any European plant grown in cultivation, but has 
assumed an "economic” role only in utilization of European plants 
(mustards) growing in semi-wild habitats. The other Pieris are 
restricted entirely to one or a few native species of plants. 

The food plant tests were made in several series at different times 
of the year and with different species of plants. Each series was run 
for the purpose of comparing the influence of several plants (usually 
five) on growth rate and mortality. For each series, a group of twenty 
larvae, obtained as follows, were grown on each plant. Female Pieris 
rapae were allowed to oviposit on a kale plant in the greenhouse. For 
the first series, the eggs were removed from the kale leaf and placed 
on leaves of the plant being tested in a petri dish. On all other tests, 
the eggs .were allowed to hatch on kale, and to eat on it for one or 
two days before removal to the tested plants. In order to minimize the 
effects of variable environmental conditions, all tests of a series were 
grown at the same time and place. For example, eggs laid on September 
11, 1959, were used for the first test. On September 14, 1959, the 
larvae were measured for the first time, and then successively thereafter 
each day until pupation. The dates of larval (or pupal) deaths were 
noted. 

The later tests were similar except that (1) the origin of the 
parents was different, ( 2 ) one or two day old larvae were used instead 
of eggs, ( 3 ) the time of year and ( 4 ) place of breeding was different. 
Data on origin of the material for each test are indicated in Table 1. 

TABLE 1 : Source of larvae used in these experiments. 


Series 

Species 

Food Source 

Locaiity 
( coilected 

Date Generations 

of experient from wild 




Western 



1 

Pieris rapae 

Cabbage 

Orange 

Sept.11,1959 

1 




County 



2 




Oct. 9,1959 

2 

3 




Nov. 25, 1959 

4 

4 




Feb. 20, 1960 

6 

5 




Feb. 20, 1960 

6 

6 




Aug. 2, I960 

11 

7 




Sept. 19, I960 

12 

8 

Pieris protodice 

Mustard 

Laguna Beach 

Aug. 27, 1959 

1 

9 


II 

San Fernando 






Valley 

July 18, 1960 

1 

10 


Cleome 

Owens Valley 

July 21,1960 

1 


i(t):2i-4S, 1962 


EFFECT OF FOOD PLANTS 


23 


SERIES NO. 1 

Series No. 1 was conducted for the purpose of comparing the 
effect of the following plants on growth rate, size and mortality: black 
mustard (Brassica nigra), garden nasturtium (Tropaeoum majus) , 
bladder pod (Isomeris arborea) and watercress {Nasturtium officinale) 
( Table 2 ) . The females used for the source of larvae were obtained 
from an open cabbage field isolated from any other cruciferous food 
plant source and over which the adults were swarming in huge num- 
bers. The probability that they came from any other source is 
infinitesimal. 

TABLE 2. Viability and growth rate of Pieris rapae grown on 
different kinds of food plants ( Series 1 ) . 


Plant 

Number 

Number and percent 
of larvae died 

Days from 
egg to 
pupation 

Range of 
variation in 
larval growth 

Nasturtium 

20 

16 

30 % 

20 

3 

Kale 

20 

2 

10% 

12 

2 

Mustard 

20 

8 

40% 

13 

3 

Watercress 

20 

18 

90% 

17 

1 

Nasturtium 

20 

16 

80% 

18 

5 

Isomeris 

20 

10 

50% 

20 

3 


The mortality (Table 2 . Series 1) was highest on watercress 
(90%), next highest on garden nasturtium (80%), then Isomeris 
(50%), mustard (40%) and least on kale (10%). The rate of 
development as determined by the first larva to pupate, was most rapid 
on kale (12 days), least rapid on Isomeris (20 days), and increasingly 
more rapid on nasturtium (18 days) , watercress (17 days) and mustard 
(13 days). The range in variation of larval growth was greatest for 
nasmrtium, since five days elapsed between the first larva to pupate 
and the last. The range as indicated in the table should be interpreted 
with the fact in mind that only a fraction of the original twenty larvae 
remained at the time of pupation. In other words, only two larvae 
pupated out of twenty that were fed on watercress. Had more survived, 
the range between the dates of pupation might have been greater. 

The growth rates of the larvae on the various plants were obtained 
by measuring ten larvae of each group each day and obtaining the 
average length. These were then plotted and a curve drawn for each. 
These curves are shown in Figure 1. 

Measurements of the larvae were started on the third day after 
hatching as they were so small prior to that time. However, great 

■^It should be noted that the watercress used in these experiments was obtained by purchase in a 
food market while all other plants were grown in the laboratory. It has occassionally been 
found that despite repeated washings, insecticide residues may be present upon such commercial 
vegetables and that these may cause death of larvae feeding upon them. 


24 


HOVANITZ AND CHANG 


I. Res. Lepid. 


differences already were apparent between the larvae grown on the 
various plants. Those on kale were 4.7mm long as compared with those 
on Isomeris which were only 1.5mm long. Those on nasturtium were 
1.8mm, those on watercress were 1.9mm and those on mustard were 
3.9mm. The kale-group larvae were over three times the size of the 
Isomeris group larvae. Increase in size of the larvae on subsequent 
days was fastest v/hen the larvae were grown on kale, second fastest on 
mustard, and about equally as poor on each of the other three plants. 
The largest sized larva was that grown on Isomeris (24mm) as com- 
pared with that on kale (22.2mm) but after eight more days of larval 
life. Those on watercress and nasturtium pupated at 20mm and those 
on mustard at 17.9mm. 

It appears that the greatest proportional size difference in the 
larval groups occurred during the first three days, but that most of the 
loss in size was later recovered by a longer growth period. 

SERIES NO. 2 

The tests of Series No. 2 were carried out in order to correct what 
were thought to be defects in the experiments of Series No. 1. These 
were two in number. The first was the high mortality in the eggs when 
they were removed from kale and placed on the other plants. This 
problem was averted by letting the larvae hatch on the kale plant before 
moving to the tested plants. The second presumed defect was the 
possibility of increased mortality of the very young larvae on the new 
plants, due solely to physical characteristics, toughness, etc. This was 
partially averted by allowing the larvae two days of feeding on kale 
before transference to the tested plants. 

One change in food plant was made in this test: radish obtained 
from a food market was employed in place of watercress. Another 
inevitable change was the climatic condition under which the tests 
were conducted. They were changed slightly by the fact that the 
tests were carried out starting October 9, 1959, instead of 

September 11, 1959. 

TABLE 3. Viability and growth rate of Pieris rapae grown 
on different kinds of food plants (Series 2). 

Days Range of 

from variation 
egg to in 

No. and % No. and % Total No. pupa- larval 

Plant No. of larvae died of pupae died and % died tion growth 


Kale 

20 

2 

10% 

0 

0 

2 

10% 

11 

2 

Mustard 

20 

2 

10% 

3 

15% 

5 

25% 

13 

4 

Radish 

20 

8 

40% 

5 

25% 

13 

65% 

13 

5 

Nasturtium 

20 

8 

40% 

2 

10% 

10 

50% 

14 

5 

Isomeris 

20 

12 

60% 

3 

15% 

15 

75% 

15 

3 


The mortality (Table 3) in this series was generallly not so high 


PIER IS RAPAE 



Growth rates of larvae of Pieris rapae grown on various plants ( Series No. 1 ) . 


26 


HOVANITZ AND CHANG 


/. Res. Lepid. 


as in the series just preceding, though that on kale was the same at 
10%, those on mustard dropped from 40% to 10%, on nasturtium 
dropped from 80% to 40% and on Isomeris increased from 50% to 
60%. Watercress was not tested this time but instead radish^ was used, 
with a resulting mortality of 40%. The better survival of these larvae 
is probably correlated with the fact that they were all grown during the 
first two days on kale. 

Generally, the time (minimum days) to pupation in this series 
has been decreased (Fig. 2), The time on kale has been decreased 
one day, on mustard it remains unchanged, and on nastrutium and 
Isomeris it has been reduced five days. These reductions are probably 
correlated with the more successful survival of the young larvae on 
kale for the first two days, as compared with Series No. 1. 

SERIES NO. 3 

The tests of Series No. 3 are identical with those of Series No. 2 
with the exception that they were started November 25, 1959 instead 
of October 9, 1959; in addition, the larv.ae by this time were four 
generations removed from the wild and inbred. 

The larval mortality for all plants other than nasturtium was in- 
creased considerably in Series No. 3 compared with Series No. 2; 
10% to 30% for kale, 10% to 35% for mustard, 40% to 80% for 
radish and 60% to 75% for Isomeris (Table 4). The data are com- 
parable for the percentage of pupae died. 

TABLE 4. Viability and growth rate of Pieris rapae grown 
on different kinds of food plants ( Series 3 ) . 

Days Range of 
from variation 


Plant 

No. 

No. and % 
of larvae died 

No. and % 
of pupae died 

Total No. 
and % died 

egg to 
pupa- 
tion 

in 

larval 

growth 

Kale 

20 

6 

30% 

0 

0 

6 30% 

15 

2 

Mustard 

20 

7 

35% 

0 

0 

7 35% 

16 

5 

Radish 

20 

16 

80% 

1 

5% 

17 85% 

19 

4 

Nasturtium 

20 

7 

35% 

1 

5% 

8 40% 

16 

6 

Isomeris 

20 

15 

75% 

2 

10% 

17 85% 

18 

8 


The length of larval life has also been increased considerably in 
Series No. 3 tests, and thus also the range of variation in larval growth. 

The reasons for these differences between Series No. 2 and No. 3 
are probably two fold. First, the later time in the season has meant 
that the conditions of growth were somewhat different, primarily in 
the temperature. 

The larvae were grown in a greenhouse in which the temperature 
was not well-regulated. Therefore, the temperature of the October 

^The radish used in this particular experiment, but not in others reported on by us, was obtained 
in a food market. Refer to footnote (4) for possible latent effects. 


(mm) 3VAMV1 JO H10N31 


(i):2i-42, 1962 


EFFECT OF FOOD PLANTS 


27 


tests (Series No. 2) was higher than for the November tests (Series 
No. 3). This would profoundly influence the results insofar as the 
absolute growth rates are concerned. On the other hand, it did not 
affect the relative growth rates on the various food plants. These are 
still in the same descending order: kale, mustard, radish, nasturtium 
and Isomeris. 



0 ■ I 1 ■ ! I I ■ ■ U— I I 

U 2x 3 4 5 m 6 r 8 9x )0 ItxK !2 13 14 15 

DAYS AFTER HATCHING FROM EGG UNTIL PUPATION NO OBSERVATION 

MX = NO OBSERVATION ON KALE 

2. Growth rates of larvae of Fieris rapae grown on various plants (Series 

No. 2). 

The differential increase in size of the larvae on the various food 
plants is better shown in Series No. 3 (Fig. 3) than in Series No. 2 
( Fig. 2 ) . This is because the slower development rate served to 
exaggerate the differences between each strain. The larvae grown on 
kale grew larger at a faster rate than the larvae on any other plant, 
followed by mustard, nasturtium, Isomeris and radish. In Series No. 2, 
the order of the latter two were reversed. It is interesting to note that 
the ultimate size of the larvae which grew slowest was seldom equal 
to the size of those larvae which grew fastest. In fact, there is almost 
a direct relationship between these events. 

A second reason for the difference between the mortality and 
growth data of these two series may be of genetic significance. The 
extent of inbreeding in the laboratory population by the fourth genera- 
tion would have been sufiicient to (a) increase the mortality, (b) slow 


20 0 1_ P/ERIS RAPAE 


HOVANITZ AND CHANG 


J. Res. Lepid. 


28 



O 


(u/m) 3VAi/Vl JO H19N31 


DAYS AFTER HATCHING FROM EGG UNTIL PUPATION (x=NO OBSERVATION) 


( i ):2 i -42, 19^2 


EFFECT OF FOOD PLANTS 


29 


development rate, (c) decrease size, (d) magnify the difficulties of 
feeding on abnormal plants by the simple means of decreasing overall 
vigor by the accumulation of allelic recessive lethal genes in the 
laboratory strain which had not yet had a chance to be eliminated by 
the selective effects of homozygosity. 

SERIES NO. 4 AND 5 

Series No. 4 was a test to compare various garden vegetables for 
their usefulness to Pieris rapae. These vegetables were mustard, kale, 
kohlrabi, broccoli and brussels sprouts. Series No. 5 was a similar test 
for various other plants. These were mustard (as before), nasturtium, 
radish, turnip and Isomeris. The tests were made in two series on 
February 20, 1962, using larvae following six generations of inbreeding, 
but of the same strain as used heretofore. In both of these tests, the 
larval mortality was higher on comparable plants used in previous tests. 
This again was probably due to the effects of inbreeding. 

TABLE 5 . Viability and growth rate of Pieris rapae grown 
on different kinds of food plants (Series 4). 

Days Range of 
from variation 


Plant 

No. 

No. and % 
of larvae died 

No. and % 
of pupae died 

Total No. 
and % died 

egg to 
pupa- 
tion 

m 

larval 

growth 

Mustard 

20 

9 

45 % 

0 

0 

9 

45 % 

15 

9 

Kale 

20 

5 

25 % 

2 

10% 

7 

35 % 

15 

10 

Kohl Rabi 

20 

9 

45 % 

0 

0 

9 

45 % 

14 

9 

Broccoli 

20 

9 

45 % 

0 

0 

9 

45 % 

14 

9 

Brussels 

Sprouts 

20 

10 

50 % 

0 

0 

10 

50% 

14 

9 


In Series No. 4 (Table 5), the data show that the larvae on kale 
had the lowest mortality rate (25%) while all the other plant tests 
were about the same (45-50%). Despite this, however, larvae from 
kohlrabi, broccoli and brussels sprouts reached pupation ahead of those 
on kale or mustard. 


TABLE 6. Viability and growth rate of Pieris rapae grown 
on different kinds of food plants ( Series 5 ) . 


Plant 

No. 

No. and % 
of larvae died 

No. and % 

of pupae died 

Total No. 
and % died 

Days 
from 
egg to 
pupa- 
tion 

Range of 
variation 
in 

larval 

growth 

Mustard 

20 

9 45% 

0 

0 

9 

45% 

15 

9 

Nasturtium 

20 

19 95% 

0 

0 

19 

95% 

16 


Radish 

20 

14 70% 

2 

10% 

16 

80% 

15 

13 

Turnip 

20 

13 65% 

0 

0 

13 

65% 

15 

4 

Isomeris 

20 

12 60% 

0 

0 

12 

60% 

18 

14 


30 


HOVANITZ AND CHANG 


/. Res. tepid. 


In Series No. 5 (Table 6), the data show that the lowest mortality 
again was on mustard but with considerably higher mortality on the 
other plants: for example, 95% on nasturtium, 70% on radish and 
60-65% on turnip and Isomeris. The length of larval life was longest 
on Isomeris (18 days) and about the same (15-16 days) on other 
plants. The range of larval growth period varied greater with radish 
and Isomeris than with any of the other. 



4 5 6 7 ex 9 10 II 12 13 14 I5x . 16 

DArS AFTER HATCHING FROM EGG UNTIL PUPATION tx^NO OBSERVATION) 


4. Growth rates of larvae of Pieris rapae grown on various plants (Series 
No. 4). 

The rate of increase in size (Fig. 4) was more rapid with broccoli, 
kohlrabi and brussels sprouts than with kale and mustard. It is believed 
that this may be correlated with a more turgid or water-content condi- 
tion of these plants as compared with the drier or stiffer condition of 
the kale and mustard. For some reason, in Series No. 5, also, the 
increase in size of the larvae was not up to expectations on the mustard 
since the larvae on radish, turnip and nasturtium all exceeded those on 
mustard in final size ( Fig. 5 ) . 

SERIES NO. 6 

Series No. 6 was tested on August 2, I960 with the use of larvae 
eleven generations from the wild. One new plant, {Cleome lutea) , was 
tested in this series, since this plant was found to be primary food 
plant for Pieris protodice in Owens Valley, California. Mortality data 


PiERiS RAPAE 


i(i) 121-42, 1^62 


EFFECT OF FOOD PLANTS 


31 




o o 00 

in o in cJ 


q 

o 

CVJ 


(u/m) BVAyVl JO H10N31 


Growth rates of larvae of Fieris rapae grown on various plants ( Series No. 5 ) . 


32 


HOVANITZ AND CHANG 


/. Res. Lepid. 


for kale , mustard, nasturtium and Isomeris, (Table 7) compared well 
with previous data obtained on these plants (Series 1, 2, 3, 4 and 5). 
Cleo7ne was only slightly less effective in maintenance of larval life, 
the mortality rate being 30% as compared with 20% for kale and 
mustard. The relative lengths of larval life and the range of variation 
in larval growth also compares favorably with previous data. The 
higher mortality as seen in Series 3, 4, 5 and 6 has here been reduced 
to nearly the level of Series No. 1 . 


TABLE 7. Viability and growth rate of Pieris rapae grown 
on different kinds of food plants (Series 6). 


Plant 

No. 

No. and % 
of larvae died 

Days from egg 
to pupation 

Range of 
variation in 
larval growth 

Kale 

20 

4 

20% 

12 

3 

Mustard 

20 

4 

20% 

13 

3 

Nasturtium 

20 

10 

50% 

13 

3 

Cleome 

20 

6 

30% 

14 

5 

Isomeris 

20 

12 

60% 

18 

6 


The differences between growth curves for each of the plants 
tested has been greatly increased in this test as compared with earlier 
tests. The factors responsible for this are unknown but may be due 
to a reduced adaptability caused by increased homozygosis (Fig. 6). 

The larvae on kale were larger than those of any other plant at all 
times during their growth period. Pupation occurred the twelfth day. 
The larvae grown on nasturtium were almost the same size as those 
grown on mustard, whereas in Series No. 1 (Fig. 1) the larvae on 
mustard grew faster and larger. The larvae on Cleome were fourth in 
size on any day, while those on Isomeris were fifth. 

SERIES NO. 7 

This series was conducted in order to place the plant Thelypodium 
affine into position relative to the other standard test plants in the 
laboratory. Thely podium affine is a native cruciferous plant from the 
desert regions which is not known to be a food plant of any Pieris 
species, though another Thely podium species is the food plant of Pieris 
dsymhrii. 

TABLE 8. Viability and growth rate of Pieris rapae grown 
on different kinds of food plants ( Series 7 ) . 


Plant 

No. 

No. and % 
of larvae died 

Days from egg 
to pupation 

Range of 
variation in 
larval growth 

Kale 

20 

5 

25% 

16 

3 

Mustard 

20 

6 

30% 

17 

4 

T. affine 

20 

14 

70% 

22 

5 

Isomeris 

20 

12 

60% 

18 

6 

Nasturtium 

29 

8 

40% 

17 

6 


PIER IS RAPAE 


■{1)121-42, 1962 


EFFECT OF FOOD PLANTS 


33 


\ 



o 

o 

CSJ 


fluuij 3VAHW1 JO HI9N31 


DAYS AFTER HATCHING FROM EGG UNTIL PUPATION 


34 


HOVANITZ AND CHANG 


/. Res, Lepii. 


The tests of Series No. 7 (Table 8) were carried out on 
September 19, I960, using larvae after twelve generations of laboratory 
inbreeding and growing on kale. The larvae grew relatively well on 
mustard, kale and nasturtium. The effectiveness of Isomeris was as 
previously tested. Thely podium affine was the poorest of all these 
plants as far as effectively providing the larva with stimulation to feed. 

The mortality of the larvae on T, affine was 10% compared with 
60% on Isomeris, 40% on nasturtium, 30% on mustard and 25% 
on kale. The length of larva life was increased from 16 days on kale 
to 22 days on Thelypodium. The final size of the larvae was reduced 
from 22mm long on kale to only 18mm long on the Thelypodium 
larvae which survived. (Fig. 7). 

GENERAL OBSERVATIONS ON PIERIS RAPAE TESTS 

The seven series of tests on Pieris rapae larvae have indicated 
several significant points: 

( 1 ) Larvae from strains grown previously on cabbage or kale 
have a lower mortality rate, have a faster development rate, and a 
greater size when grown on kale than when grown on any other plant 
tested. When similar larvae are grown on mustard, these factors are 
decreased slightly. The order of decrease for plants other than these 
two is in the order approximately as follows: nasturtium, Cleome, 
radish, Isomeris, and T, affine. Plants of the cabbage group: kohlrabi, 
broccoli and brussels sprouts, are about the same as cabbage or kale. 
Turnip is about the same as radish. 

( 2 ) Higher mortality in the tests is generally correlated with a 
slower development rate, a delayed time to pupation and smaller size 
of the larva and pupa. There are some exceptions to this rule; for 
example, the final size of the larva may be as great when the growth 
rate was slow on one plant as when it was fast on another. This has 
been indicated, however, only for plants of the cabbage group. 

(3) Mortality rates in the various test series have been variable. 
These variations in mortality have been thought not only to be due to 
the genetic effects of inbreeding but also partly to the changed environ- 
mental conditions encountered while the tests were conducted, such as 
lower temperatures, etc. of the changing seasons. Lower temperatures 
slow the developing rate, and increase the relative humidity. Increased 
humidity permits the larvae to be attacked by bacterial or viral diseases 
to a much greater degree. 

SERIES NO. 8 

Tests of Series No. 8, 9 and 10 were conducted with larvae of 
Pieris proto dice rather than with Pieris rapae. 

The first series with this species (No. 8) was conducted on 
August 27, 1959, utilizing larvae one generation from the wild. The 



fwu/} 3VAHV1 JO H10N31 


DAYS AFTER HATCHING FROM EGG UNTIL PUPATION ( NO OBSER V ATION 


36 


HOVANITZ AND CHANG 


/. Res. Lepid. 


females of the previous generation were obtained in a field of black 
mustard, not near any other source of food plant, and in which eggs 
and larvae were found on that plant. This is good' evidence that the 
natural plant utilized here as larval food was mustard. 

In this series, the food plants tested as larval food were mustard, 
kale, radish, Isomeris and nasturtium. The mortality of larvae of 
Pieris protodice on these plants increased in the order just stated, 
namely, 10% on mustard, 20% on kale, 55% on radish, 55% on 
Isomeris and 95% on nasturtium (Table 9). Deaths in the pupa stage 
on kale, radish and Isomeris materially increased the total mortality 
before adult emergence. This was not increased on mustard. 


TABLE 9. Viability and growth rate of Pieris protodice grown on 
different kinds of food plants (Series 8). 


Plant 

No. 

No. and % 
of larvae died 

No. and % 
of pupae died 

Total No. 
and % died' 

Days 
from 
egg to 
pupa- 
tion 

Range of 
variation 
in 

larval 

growth 

Mustard 

20 

2 

10% 

0 0 

2 

10% 

10 

2 

Kale 

20 

4 

20% 

2 10% 

6 

30% 

11 

2 

Radish 

20 

11 

55% 

4 20% 

15 

75% 

13 

3 

Isomeris 

20 

11 

55% 

3 15% 

14 

70% 

16 

3 

Nasturtium 

20 

19 

95% 

■— — - 

19 

95% 

21 



The length of larval life was greatly different on the various food 
plants, ranging from ten days on mustard, eleven days on kale, thirteen 
days on radish, sixteen days on Isomeris to twenty-one days on nastur- 
tium. This wide variation in length of larval life is greatly different 
from anything encountered with Pieris rapae. It would indicate a 
fundamental difference in food plant adaptability between these two 
species, Pieris rapae being much more versatile in its requirements than 
Pieris protodice. 

The size of the larvae at various days after hatching (Fig. 8) 
indicates that the mortality data are in close correlation with the size 
increments of the larvae with respect to food plants involved. Larvae 
on mustard and kale increased to 24mm in length, those on radish, 
Isomeris and nasturtium to 20-2 1mm each. 

SERIES NO. 9 

Discovery that Pieris protodice utilizes Cleome lutea as a natural 
food plant in the Owens Valley prompted a series of tests involving 
this plant. Both Series No. 9 and Series No. 10 were carried on for 
the purpose of testing Cleome in comparison with other plants used 
as standards. Series No. 9 utilized Pieris protodice larvae bred from 
adults obtained from mustard fields in the San Fernando Valley while 
Series No. 10 utilized Pieris protodice larvae obtained from Cleome 


pf£ms pporoDicE 


i(i);2i-42, i$62 


EFFECT OF FOOD PLANTS 


37 



mrs AFTER HATCHING FROM EGG UNTIL PUPATION (NO observation = x j 


38 


HOVANITZ AND CHANG 


/. Res. Lepid. 


plants in the Owens Valley (Big Pine). There is no chance that the 
San Fernando population had access to Cleome but there is a chance 
that the Owens Valley population did have limited access to mustard. 

TABLE 10. Viability and growth rate of Pieris proto dice grown on 
different kinds of food plants (Series 9). 


Plant 

No. 

No. and % 
of larvae died 

Days from egg 
to pupation 

Range of 
variation in 
larval growth 

Mustard 

20 

6 

30% 

14 

3 

Cleome 

20 

7 

35% 

14 

4 

Nasturtium 

20 

18 

90% 

17 

2 

Isomeris 

20 

8 

40% 

17 

5 


The Series No. 9 larvae (Table 10) showed highest mortality on 
nasturtium (95%), just as in Series No. 8, with the lowest mortality 
on mustard (30%), next highest on Cleome (35%) and then Isomeris 
(40%). This corresponds well with comparable data in Series No. 8. 

The growth increment curves for this series are somewhat com- 
parable with the preceding series except for the one surviving larva 
on nasturtium which increased to a size greater than that attained by 
larvae on any other plant. The differences between the growth curves, 
however, are not as distinct and clear as in Series No. 8, perhaps because 
of temperature differences between the two. Cleome is almost as 
satisfactory as mustard as a food plant for these larvae. ( Fig. 9 ) • 

SERIES NO. 10 

This series was run almost contemporaneously with the preceding 
and involved larvae from the Owens Valley strain grown on Cleome. 
The mortality (Table 11) was lowest on mustard (20%), second 

TABLE 11. Viability and growth rate of Pieris protodice grown on 
different kinds of food plants (Series 10). 


Plant 

No. 

No. and % 
of larvae died 

Days from egg 
to pupation 

Range of 
variation in 
larval growth 

Mustard 

20 

4 

20% 

11 

3 

Cleome 

20 

6 

30% 

12 

5 

Isomeris 

20 

7 

35% 

13 

7 


lowest on Cleome (30%) and then Isomeris (35%). There was a 
greater range of larval growth rates on this group of larvae on Isomeris 
than in any other group as well as the lowest mortality on this plant 
in any of the larvae tested. The growth increment curves for these 
larvae indicate that the larvae do not show much difference in their 
reaction to mustard and Cleome but that mustard is still slightly better, 
and Isomeris is always poorer. Final size of the larvae, however, was 
larger for the Cleome (23 mm) than for the mustard (22 mm) or 
the Isomeris (18 mm)'. (Fig. 10). 


LEN6TH OF LARVAE (mm) LENGTH OF LARVAE (> 


i(i):2\-42, 1962 


EFFECT OF FOOD PLANTS 


39 



DAYS AFTER HATCHING FROM EGG UNTIL PUPATION 

9 . Growth rates of larvae of 'Pieru protodice grown on various plants (Series 
No. 9). 



PAYS AFTER HATCHING FROM EGG UNTIL PUPATION 


10. Growth rates of larvae of Pieris protodice grown on various plants (Series 
No. 10), 


40 


HOVANITZ AND CHANG 


/. Res. Lepid. 


GENERAL OBSERVATIONS ON PIERIS PROTODICE TESTS 

The three series of tests on Pieris protodice have indicated several 
significant points: (1) Larvae from strains obtained from mustard 
or Cleome populations had equally good survival value on mustard 
plants. The following plants gave decreasing survival value to the 
larvae: (Series 8) kale, radish, Isomeris and nasturtium. (Series 9) 

Cleome, nasturtiuna, Isomeris. (Series 10) Cleome, Isomeris. (2) Higher 
mortality in the tests is generally correlated with a slower development 
rate, a delayed time to pupation and smaller size of larvae and pupae. 

DISCUSSION 

The tests described in the preceding pages have indicated some 
significant information regarding the effect of various food plants on 
survival in two species of Pieris. The ability to survive on different 
food plants is not an all or none relationship, but rather is a relation- 
ship based upon relative ability to survive. In the most extreme case, 
all larvae may die rather than eat a completely unacceptable plant. 
On the other hand, plants may be placed in an order of desirability 
with regard to the ability of larvae to survive, or to grow satisfactorily, 
on plants which they may accept at least reluctantly. 

The tests described in this report are based upon using three criteria 
to indicate the desirability of a plant to the larvae. The first of these 
is the ability to survive, or the reverse which is the mortality, expressed 
as a percentage of the total larvae which died during the experiment. 
The second criterion is the daily size increment during the life of the 
larva. This actually tests two factors: (1) the physiological usefulness 
of the food to the larvae and (2) the total amount of food eaten and 
uilized, which may be a direct result of the stimulatory activity of the 
plant toward the larvae. The third criterion is the range in variation 
of larval growth. The significance of this criterion is in more doubt 
than the other two, but the range indicated would in some degree show 
the genetic variability present in the larval population toward physio- 
logical adaptation to the particular plant. 

The results just described show that some plants are much more 
suitable for larval food to certain larvae than are other plants. The 
reasons for this greater suitability are not made clear. In those cases 
where higher mortality, slower growth rate, size increments and final 
size are correlated, the cause of the poor growth could be lack of 
nutriment in the plant, or lack of larval ingestion of the plant. In other 
cases where larval mortality may be high, as with the broccoli, Kohl- 
rabi and Brussels sprouts tests (Table 5) hut growth rate rapid and 
size large ( Fig, 4 ) , there are probably two factors involved. The rapid 
increase in size could be due to a high attraction to a very nutritious 
plant, but secondary conditions surrounding the plant may be conducive 


I (i):2i -42, 1962 


EFFECT OF FOOD PLANTS 


41 


to a higher mortality, namely, high water content in the plant itself, 
or higii humidity surrounding tne plant whicn would increase the 
susceptability of the larvae to virus and bacterial intections. 

Toughness of leaves of a plant cannot be ruled out as a factor 
involved in larval speed of growth. Some leaves are too tough for the 
larvae to feed upon easily. Leaves of the same plant vary in tnis regard; 
succulent leaves at the siioot apex are usually much more desirable than 
older, mature leaves. This may be caused by toughness, or it may be 
caused by the greater perception of attractive substances permeating 
from younger leaves. At any rate, the relative uniformity of the tests 
in the difterent series is good evidence for the conclusion that this 
factor has not been of major significance. 

The greater ability of tieris rapae as compared with Pieris protodice 
to survive on a variety of common cruciferous plants is interesting in 
view of the fact that both succeed well on black mustard. Pieris rapae 
survives better on kale over mustard and P. protodice survives better on 
mustard over kale. However, on most other cruciferous plants tested, 
there was a wide disparity between their preferences. The selections 
made by the strains of larvae shown by these experiments complements 
perfectly the known differences in geographical and ecological distribu- 
tion of the two species concerned, and indicates that there is little 
natural competition between the species for food plants. This shows 
also why Pieris rapae is always associated with European cruciferous 
plants, weeds or truck crops, while Pieris protodice is generally 
associated with other cruciferous plants. 

Takata (1957, 1959, 1961) has published a number of papers 
which bear upon this problem. None of these deals precisely with 
the effect of various food plants on survival and growth rate of Pieris, 
but rather deals only with selection preferences by the larvae and the 
adult. Further considerations of the relation of this work of Takata 
on Pieris rapae crucivora to the present general program will therefore 
be made in a complementary paper entitled "Three Factors affecting 
larval choice of food plant” to be published in this journal. 

A visit to our project by W. H. Dowdeswell of England during the 
course of these experiments was followed by a similar study of more 
limited extent on Pieris napi L. in England. Dowdeswell found that 
two populations of this Pierid were feeding on different cruciferous 
plants. By switching the food plants in laboratory experiments on test- 
ing the larvae, he found as we had, that there was a higher mortaity 
and decreased growth rate when the larvae were fed on plants not 
the normal food plant for that population (Dowdeswell and Willcox, 
1961 ). 

SUMMARY 

1. Tests were made of the ability to survive of larvae of two species 
of Lepidoptera, Pieris rapae and Pieris protodice on various food plants. 


42 


HOVANITZ AND CHANG 


/. Res. tepid. 


2. Three criteria were tested: (a) mortality, (b) growth rate 
and size increment, (c) range in variation of larval growth. 

3. Seven series of tests involving Pieris rapae were conducted 
showing the results summarized under the heading "General Observa- 
tions on Pieris rapae Tests.” 

4. Three series of tests involving Pieris protodice were conducted 
showing the results summarized under the heading "General Observa- 
tions on Pieris protodice Tests.” 

5. Pieris rapae has significantly different requirements for food 
plants than Pieris protodice, even though the tested larvae were both 
from mustard-bred strains. 

6. The native adaptability of Pieris rapae to survive on various 
plants is better than that of Pieris protodice. 

7. The causes of differences in survival value of the larvae on 
different food plants are believed to be several, but the main one is 
the total amount of food eaten and utilized. 

LITERATURE CITED 

DOWDESWELL, W. H. AND H. N. A. WILLCOX. 1961. Influence of the 
food plant on growth rate and pre-imaginal mortality in the green-veined 
white butterfly Pieris napi (L). Entomologist 94:2-8. 

HOVANITZ, W. 1962. The ecological and geographical distribution of 
Pieris species in Southern California. J. Res. Lepidoptera 1 : in press. 
HOVANITZ, W. 1962. The distribution of the species of the genus Pieris 
in North America. J. Res. Lepidoptera 1 ( 1 ) : this issue. 

TAKATA, N. 1957. Studies on the host preference of cabbage butterflies 
{Pieris rapae L.) III. Mechanism of food preference of cabbage butterfly 
larvae. Jap. J. Ecol. 7 (3) :117-119. 

TAKATA, N. 1959. Studies on the host preference of common cabbage 
butterfly, Pieris rapae crucivora Boisduval. IV. Tendencies of food prefer- 
ence in relation to total quantity of the host plants cultivation. Zoological 
Magazine (Japan) 68(5) : 187-192. 

TAKATA, N. 1959. Studies on the host preference of common cabbage 
butterfly, Pieris rapae crucivora Boisduval. VI. Change in the food 
preference of larvae when reared successively by the definite food plant 
for several generation. (Preliminary report) Jap. J. Ecol. 9:224-227. 
TAKATA, N. 1961. Studies on the host preference of common cabbage 
butterfly, Pieris rapae crucivora Boisduval. XI. Continued studies on the 
oviposition preference of aduir butterflies. Jap. J. Ecol. 11 (3) :124-133. 
TAKATA, N. 1961. Studies on the host plant preference of the common 
cabbage butterfly, Pieris rapae crucivora Boisduval. XII. Successive 
rearing of the cabbage butterfly larva with certain host plants and its 
effect on the oviposition preference of the adult. Jap. J. Ecol. 11(4): 
147-154. 


Journal of Research on the Lepidopvera 


l(l):43-49, 1962 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
Copyright 1962 


GENERAL CHARACTERISTICS OF THE 
MOVEMENTS OF VANESSA CARDUI (L.) 

J. W. TILDEN 

San Jose State College, San Jose, Calif. 

It is a matter of common knowledge that the populations of 
many species of butterflies participate in large and extensive mass 
movements. While several workers, notably C. B. Williams of England, 
have done extensive work in this field, relatively few investigators 
have made it their principal subject of research. Though much 
pertinent information has been assembled and certain tentative con- 
clusions may be reached, it cannot be said that the movements of 
butterflies are as well understood as certain other phases of the study 
of these insects. 

Most movements of butterflies, while usually referred to as migra- 
tions, would not meet the criteria set up by students of vertebrates for 
defining migrations. The migrations of birds, for example, involve 
cyclic movements. There is a going out and a return. Few butterflies 
other than the Monarch [Danaus plexippus (L.)], habitually possess 
such a rhythmic pattern of movement. Even with the Monarch, the 
individuals that return are usually (probably always) not the same 
individuals that initiated the outward movement, but are the offspring 
of these original individuals. 

Many movements of butterflies are relatively local wanderings, and 
may be in one direction in one local area, but in a different direction 
in another area a relatively few miles away. For example, in the fall of 
I960, Tilden noted Libytheana hachmanii (Kirt.) (the Snout Butter- 
fly) moving southwesterly in large numbers in the vicinity of Con- 
tinental, Pima Co., Ariz., on Sept. 14. The next day, Sept. 15, equally 
large numbers of this species were observed moving up-canyon ( north- 
ward) in Sycamore Canyon, Santa Cruz Co., about thirty -five miles 
south of Continental by air line, though considerably further by road. 

The very conspicuous movements of certain species have been 
regarded as depending on population pressures, leading to a mass 
exodus. These may be considered as emigrations from the populated 
area, and as immigrations into surrounding area. Such movements 
frequently appear to be in one direction only. Unless the insects were 
to find suitable conditions at the far end of the movement, establish- 


43 


44 


TILDEN 


/. Res. Lepid. 


merit would not result. Such movements have been thought to account 
in part for the finding of strays far beyond the normal ranges of 
the species. 

Movements of an altitudinal nature have been observed. Nymphalis 
calif ornica (Bdv.) has been seen ascending the Sierra Nevada of 
California in great numbers. Such movements may result in finding 
sufficient larval food plants for a subsequent brood. In I960, Tilden 
found this species in outbreak numbers of adults at Manzanita Lake, 
Lassen Volcanic National Park, Calif., on June 10-17. Numerous larvae 
in various stages of development were found on Ceanothus spp. Adults 
could be observed many miles below the Park, working their way up 
the mountain. It may be significant to note that this species passes 
the winter as an adult. It seems plausible that these altitudinal move- 
ments of adults are the usual way for adults to reach the higher eleva- 
tions, where they act as parents of a late high elevational brood. This 
might be a fruitful field of investigation. 

The often spectacular "migrations” of Vanessa cardui (L.) present 
an interesting condition somewhat intermediate between true migra- 
tions and other types of movements. Return movements seem not to 
be clearly documented in regard to this species. However, reproduction 
has been observed repeatedly along the route, and the offspring take 
up the movement when they emerge as adults. These mass movements 
of Vanessa cardui thus proceed in periodic waves, alternating with 
relative lulls between, and the whole process has much more continuity 
than is characteristic of the movements of most other butterfly species. 

Much of the basic work on the movements of Vaness-a cardui has 
been done by C. H. Abbott. Others, including Sudgen, Woodbury and 
Gillette in Utah, have made contributions. Among the various titles 
that form the literature on the movements of this species in western 
United States, some are short observational notes only. Others present 
a considerable amount of detailed study. Abbott has attempted to 
understand the nature of these movements and has produced several 
penetrating papers on the subject. 

The last great movement was in 1958. Abbott traces three or 
more generations extending up to or beyond the Central Valley of 
California. Other generations carried the waves through the Bay 
Region of California on into Oregon. The generations involved in 
the northern portions of the movement become less clear. 

The first generation seems to have originated in northern Mexico, 
the second in the Imperial Valley of California, western Arizona and 
the eastern Mohave Desert of California, the third in western Mohave 
Desert, and the fourth in the Central and Coastal Valleys of California, 
and to the Bay Region. Abbott (1959) has presented material that 
includes the observations of a number of workers, and has coordinated 
this material. In addition, the follov/ing observations are here presented 
for the first time. 


j(i): 4 }- 49 , 1962 


MOVEMENTS OF VANESSA 


45 


Tilden found many dead adults in western Arizona (Mohave 
County) and in the vicinity of Needles, California, as well as large 
numbers of moving adults, March 30-31, 1958. From Barstow, San 
Bernardino Co., west to Mohave, Kern Co., great numbers of adults 
were flying northwesterly on March 31. Later that day adults were 
seen moving through Tehachapi Pass, Kern Co., and spilling into the 
San Joaquin Valley east of Bakersfield, Kern Co. On April 20 to 
May 9, eggs and larvae were abundant in the Salinas Valley, 
Monterey Co. 

On May 30 - June 1, Tilden found large numbers of adults along 
the Redwood Highway (Route 101) between San Rafael, Marin 
County, and Cloverdale, Sonoma County, moving northwesterly. Dead 
adults were seen as far west as Booneville, Mendocino County, where 
but a few living adults remained. This may mark the western edge 
of the movement. 

In the vicinity of The Geysers, Sonoma County, adults were moving 
northwesterly in large numbers in overcast weather (fog) on June 1, 
before noon. Larvae were abundant on Amsinckia (Fiddleneck) and 
on the introduced Spanish Thistle ( Carduus ) , in various stages of 
development. 

Two interesting reports were from Oregon. Ray Albright reported 
large numbers of Vanessa cardui moving in a north-north-easterly 
direction on May 18-19, 1958, while on May 20-21, equally large 
numbers were observed moving in exactly the opposite direction. He 
attributes this reversal of direction to the insects meeting a cold front, 
since a storm was in progress some distance beyond the place toward 
which the insects were noted to be flying. The other report was from 
David Huntzinger at Crater Lake National Park. He noted that Vanessa 
cardui was abundant there in 1958, though scarce or absent in 1957. 

On April 1, 1958, Tilden observed what is apparently a new 
observation for the United States. Large numbers of Vanessa cardui 
were flying north-westerly in the rain. Between Vidal, San Bernardino 
County, and Desert Center, Riverside County, the rain was light to 
moderate, but rather cold. Between Indio and Whitewater Canyon, 
later the same day, the flight still continued in heavy to very heavy 
rainfall. This seems to indicate that adults may continue to fly, once on 
the wing, even in heavy precipitation. It would seem a priori that 
initiation of a flight during rain might be less likely. 

From observations so far recorded it is possible to generalize on 
some of the characteristics of the movements of Vanessa cardui. Some 
of these generalizations have been set forth by Abbott and others, 
while some are suggested here for the first time. 

1. The adults tend to fly into the wind. This may be a key point 
in attempting to explain the prevailing direction of the movements. 
During the time of year when Vanessa cardui is moving, the prevailing 
winds in California are north-westerly, and the movements of the 


46 


TILDEN 


/, Res. Lepid. 


butterflies are also in a north-westerly direction, almost or quite directly 
into the prevailing winds. Sudgen, in 1937, noted a northerly direction 
of flight in Utah. Sudgen, Woodbury and Gillette, in 1947, noted a 
north-easterly direction of flight. Observations to be made in the future 
might very well take special note of the wind direction as well as the 
direction of the insects’ flight, to find how complete this correlation 
between wind direction and flight direction may be. 

2. The adults move at a rather equal velocity in relation to one 
another, resulting in a fairly constant spacing of the individuals. 
There have been some observations made where this did not seem to 
be the case, but for the most part, this generality seems to hold. 

3. The insects tend to fly over obstacles rather than around them. 
At times this appears to the observer quite amusing, even ridiculous. 
If a tree, building or parked car obstructs the route, the adults fly over 
the top of the obstruction, when in some instances it would appear 
that a veering to the right or left would be more easily accomplished. 
The obstruction, whether it be low or tall, is barely cleared. On 
March 30, 1958, Tilden observed them piling directly over a small 
building near Barstow, San Bernardino County, the only building for 
miles in an open desert. 

4. The adults tend to fly at rather even densities. If a line is 
marked off between tv^o points, fifty or one hundred feet apart, and the 
insects crossing this line per minute are counted, the results from 
several one minute counts are surprisingly similar. If then, the observer 
drives his car several miles, sets up another station and makes another 
series of counts, the results will again be similar, and frequently quite 
close to those of the previous station. But in widely separated areas, 
the count may be very different. That is, while the density on the 
Mohave Desert, for instance, may be quite constant, counts made in 
San Diego County at another time might indicate a different density. 
Further study may shed light on the possibility that density of flight 
is related to density of population. 

5. Vanessa cardui adults will fly in overcast, at least after the 
movement is under way. This has been observed between Victorville 
and Needles, San Bernardino County, March 31, 1958, and near The 
Geysers, Sonoma County, May 31 and June 1, 1958. 

6. They have been observed once at least in the United States 
(vide supra) to fly in large numbers during moderate to very heavy 
rainfall. 

7. The migrating individuals are all of one species. Other species 
of butterflies do not seem to become involved in the mass movements 
of Vanessa cardtii. 

8. The individuals in the flights are of both sexes. This was noted 
independently by sampling in 1958, by both Tilden and O. E. Sette. 
The sex ratio of the insects when engaged in a flight is very nearly 
the same as that oi Vanessa cardui at any other time, 0.5 or one half 


i ( i ):43-49> 1962 


MOVEMENTS OF VANESSA 


47 


of each sex. Tilden took a sample of fifty adults, and found twenty- 
seven males, twenty-three females. The numbers taken by Sette are 
not recorded. This phase of the investigation deserves further study 
and this is planned for the future. 

9 . The females from the above sample were not greatly gravid. 
None had eggs ready to oviposit. This suggests that the females may 
drop out of the flight when ready to oviposit. If this be true, it might 
explain the tendency to reproduce along the flight routes. 

10. The total movement proceeds in waves, or phases, which so 
far appear to be based on broods, a conclusion which seems to be well 
supported by the work of Abbott. 

In the light of current findings, it is possible to reconstruct, at 
least to some degree, the nature of the populations of Vanessa cardui 
during and prior to the flights. 

1. The first (initiating) brood apparently originates to the south, 
in northern Mexico or southwestern Arizona. 

2. Findings from those movements that have been studied indicate 
that large flights can occur only in years when there has been sufficient 
rainfall on the deserts for a large growth of vegetation, enough to 
support such populations of Vanessa cardui. 

3 . The food plants in the desert areas, as noted by many observers, 
are principally Boraginaceae, especially Cryptantha, but also Amsinckia. 
Malvaceae have also^ been reported. Thistles and other composites seem 
to be of little importance at this time. 

4. Upon emergence of the adults, each succeeding brood begins 
to fly into the wind, which in the deserts of California at this season 
(early spring) results in a north-westerly direction of flight. 

5. These adults reproduce at some point along the line of flight. 
Time and place of reproduction presumably depend on the condition 
of the insects and the presence of suitable vegetation. Information on 
how far the females fly before reproducing, and the precise reason 
for selecting a certain place for reproduction, would add much to our 
knowledge. 

6. Four to five broods, perhaps more, succeed one another before 
the force of the movement is spent. 

7. The northern extent of the movement is at least to the San 
Francisco Bay Region of California. In some years, such as 1958, the 
effect extends into northern California, and apparently further, to 
Oregon and perhaps even Washington. Information as to the relation- 
ship between the Vanessa cardui in California and those in Oregon 
and Washington during such outbreak years is badly needed. 

Williams and Abbott have raised the question as to whether 
Vanessa cardui is found north of the Imperial Valley, Imperial County, 
Calif., in any but outbreak years. There seems to be good evidence 
that it is. An examination of pinned specimens from many parts of 
California shows specimens taken in almost every year for many years 


48 


TILDEN 


/. Res. Lepid. 


back. As a Professor at San Jose State College, Tilden has examined 
hundreds of student collections over a period of fourteen years, and 
has also examined other specimens dating back to 1920. Some speci- 
mens of Vanessa cardui appear in student collections every semester. 
It is evident that this species is a normal component of the butterfly 
fauna in many parts of California. It is probable, however, that this 
local endemic population has little or nothing to do with the mass 
movements which from time to time are superimposed on it. 

Two other items seem worth mentioning. Firstly, moderate to 
large populations of Vanessa cardui may be found late in the fall at 
high elevations in the mountains of the western United States. Specifi- 
cally, Tilden found hundreds of adults at Hannagan’s Meadows, White 
Mountains, Arizona, September 12, I960. A fair population was 
found at Barton Flats, San Bernardino County, Calif., in September, 
1957. Similar populations have been reported from the Sierra Nevada 
of California and elsewhere. The nature of these populations is not 
known, but it would seem that they are independent of the populations 
concerned with the great flights that occur in certain years. 

Secondly, the mass flights of Vanessa cardui are of a very irregular 
nature. They are not regularly cyclic and do not occur at predictable 
intervals. However, some success in predicting these flights has been 
possible through knowledge of rainfall on the desert. Several workers 
foresaw the 1958 outbreak in the gathering populations of the south- 
west deserts. 

An interesting and provocative suggestion of how rainfall may be 
necessary for mass movements of this species, was observed by Tilden 
September 11, I960, at Yucca, Mohave County, Arizona. Late summer 
rains were plentiful and vegetation was well developed. Numerous 
freshly emerged adults of Vanessa cardui were present, and reproduc- 
tion was in progress, with larvae of various stages on Cryptantha. It 
seemed that the basis for a movement was all prepared. However, the 
winter rains did not materialize, and on March 26, 1961, the area 
around Yucca was dry and no Vanessa cardui were to be found. If 
this abortive season is contrasted with the abundant rainfall of 1958 
over the entire area, it points to a dependence of these populations 
on ample winter rainfall in the areas that are far enough south to act 
as reservoirs for populations that can engage in a mass flight. 

Several questions are raised : ( 1 ) Why do these insects move at 

all, rather than remaining where they are? Possibly food may become 
^scarce, but populations exist that do not move out in such a systematic 
manner under lowered food conditions. (2) Why is the flight directed? 
Why do not these insects move randomly in any direction open to 
them? The tendency to fly into the wind may be a clue here. Or have 
such movements been repeated so many times over the history of the 
species, that the tendency to fly in one direction has been selected for? 
(3) What causes the adults to reproduce here and there along the 


(i):43-49< ^9^2 


MOVEMENTS OF VANESSA 


49 


route? Why do they not merely fly on until exhausted? Present 
information may suggest some tentative answers, but such answers, 
while plausible, are not yet supported by suflicient data. 

The author would add that this attempt at a synthesis of our 
knowledge concerning the movements of Vanessa cardui in western 
United States is prompted more by interest on his own part than by 
a desire to inform others. The purpose of this paper is to bring into 
focus our ignorance of the real basis of this interesting phenomenon, 
with a hope that future work will help to clarify some of the little 
known facets of the problem. 

LITERATURE CITED 


ABBOTT, C. H. 1941. The 1941 migration of the painted lady butterfly, 
{Vanessa cardui) in southern California. Bull. Ecol. Soc. Amer., 22:13. 

— — — 1946. Mapping the 1945 migration of Vanessa cardui in southern 

California. Bull. Ecol. Soc. Amer. 27 :49. 

— — 1950. Twenty-five years of migration of the painted lady butterfly, 

Vanessa cardui, in southern California. Pan-Pac. Ent., 26:616-172. 

— - 1951. A quantitative study of the migrations of the painted lady 

butterfly, Vanessa cardui L. Ecol. 32:155-171. 

— — 1959. The 1958 migration of the Painted Lady Butterfly, Vanessa 

cardui (Linnaeus), in California. Pan-Pac. Ent., 35:83-84. 

ESSIG, O. E. 1926. A butterfly migration. Pan-Pac. Ent., 2:211-212. 

SMITH, RAY F., & E. GORTON LINSLEY. 1945. Migration of Vanessa 
cardui (Linn.). Pan-Pac. Ent., 21:109. 

SUGDEN, JOHN W. 1937. Notes on the migrational flights of Vanessa 
cardui in Utah. Pan-Pac. Ent., 13:109-110. 

SUGDEN, JOHN W., ANGUS M. WOODBURY & CLYDE GILLETTE. 
1947. Notes on the migration of the painted lady butterfly in 1945. 
Pan-Pac. Ent., 23:79-83. 

WILLIAMS, C. B. 1958. Insect Migration, xiii -f235 pp., 11 plates, 22 
photos, 49 maps & diagrams. Collins, London. 

WOODBURY, ANGUS M., JOHN W. SUGDEN & CLYDE GILLETTE, 
1941. Notes on migrations of the painted lady butterfly in 1941. Pan-Pac. 
Ent., 18:165-176. 



Journal of Research on the Lepidoptera 


1 ( 1 ) : 51 - 61 , 1962 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
Copyright 1962 


THREE FACTORS AFFECTING 
LARVAL CHOICE OF FOOD PLANT’ 

WILLIAM HOVANITZ AND VINCENT C. S. CHANG" 


California Arboretum Foundation, Inc., Arcadia, 
Los Angeles State College, Los Angeles and 
University of California, Riverside 


Prior to the initiation of comprehensive tests of the attraction 
by larvae of Pieris rapae of various food plants, a number of prelim- 
inary trials were made to test the effect of external factors which might 
influence the correctness of these tests. Of the factors which were 
thought most important, the following were selected for trial: 

( 1 ) Influence of direction of air movement ( "wind”) . 

(2) Influence of age (size) of larvae. 

(3) Influence of food previously eaten by larvae, or strain of 
larvae. 


EFFECT OF WIND MOVEMENT 

An experiment devised to test this factor was set up as follows: 
A nursery flat about 24 inches square was filled with vermiculite. Pots 
of plants, each about the same size, were placed in the vermiculite 
spaced about as shown in Figure 1. The table under the flat was 
marked A, B, C and D. The direction of the wind movement was at 
side B. In the tests, larvae were placed in the central area, shown by a 
small circle and allowed to go to the plants, or to leave the flat. 

Ten different larvae were used for each experiment, and these were 
used a total of ten times each. Thus, each experiment involved 100 
trials. 

The wind direction relative to- the position of the plants was 

changed by turning the flat by 90° after each 100 trials, as shown by 
the illustration (Fig. 1). The first position tested was with mustard 
toward the dirction from which the breeze came ( side B ) . The results 
of the preferential selection by the larvae are shown on Table 1. 

Kale was selected by 19%- of the larvae, mustards by 56%, radish 

1 Aided by a grant from the National Science Foundation, Washington, D.C. 

2The authors wish to express their great appreciation to Dr. W. S. Stewart, Director, Los Angeles 
State and County Arboretum, for great cooperation in making this work possible. 


51 


52 


HOVANITZ AND CHANG 


/. Res. Lepid. 


TABLE 1. The effect of wind movement on the choice of food 
plants by larvae of Pieris rapae 



Kale 

Mustard 

Radish 

Nasturtium 

Isomeris 

None 

Total 
no. of 
trials 


A 19 

B56 

C18 

D3 

D 1 

3 

100 

90° 

D20 

A 20 

B52 

C4 

C2 

2 

100 

90° 

C43 

D25 

A 26 

B2 

BO 

4 

100 

90° 

B53 

C 15 

D13 

A8 

A5 

6 

100 


Ten 14.1 mm to 15.7 mm kale-bred larvae of Pieris rapae were used per test 
with each test ten times at each position. The parents of the larva were collected 
at western Orange County in a cabbage field. 


by 18%, nasturtium by 3% and Isomeris by 1%. Larvae which left 
the flat without selection amounted to the additional 3%. 

With the flat turned 90” so that the radish was now in position B 
( in the direction of the wind ) , the selection results were very different. 
Kale remained relatively unchanged at 20%, mustard dropped from 



(i); 5 i~ 6 i, 1962 


THREE FACTORS 


53 


56 % to 20%, radish rose from 18% to 52% at position B, while the 
remainder were unchanged. 

With a further 90° turn in the direction of the arrows (Fig. 1), 
the radish plant was now opposite the direction of the wind, and 
nasurtium-Isomeris was toward the wind. Kale at position C to the 
right side now rose from 20% to 43%, mustard at position D (away 
from the wind) rose slightly and v/as now 25%, radish at the left side 
was now 26% and nasturtium-Isomeris (toward the wind) was still 
relatively unchanged at 2 % . 

The final 90° turn placed kale in the position toward the wind. 
Here, 53% of the larvae went to kale, 15% to mustard (right side), 
13 % to radish (opposite side), and 13% to nasturtium-Isomeris 

(left side) . 

CONCLUSIONS: 

( 1 ) Of the four sides of the flat which could face the direction 
of the wind, the plants on that side were favored in three of the four 
arrangements, namely, mustard, radish and kale. Only when the 
nasturtium-Isomeris combination v/as toward the wind was the fre- 
quency in position B not increased. In fact, at that time, it was 
decreased. 

(2) These tests, of course, cannot indicate which of the three 
favored plants are relatively more favored since they each received 
about the same percentage of selection when toward the wind. However, 
when away from the wind, radish received fewer selections than either 
kale or mustard. 

(3) The Isomeris-nasturtium combination received its highest 
selection when in position A (left side). In this position also, more 
larvae left the flat without a plant selection than in any other position. 

(4) The higher selection of kale (43%) while at position C 
(right side), than radish (26%) while at position A (left side) seems 
to indicate a preference for kale rather than for radish. 

( 5 ) The lowest selection for the nasturtium-Isomeris combination 
was when it was in position B (toward the wind). This would seem 
to indicate that these plants (and Isomeris especially) repel rather 
than attract larvae of Pieris rapae. 

(6) The data indicate that the direction of wind movement is of 
great importance in larval selection of food plant, since in all cases 
where the plant has a positive effect on selection, that position in the 
direction of the wind has by far the greatest selective influence. It 
is apparent that the reverse is also true, that if a plant is repellent, 
the effect is greatest when the wind comes from that direction. 

[NOTE: The differences between the figures indicated are so great that 
the element of chance being involved is negligible. A chi-square test on the 
first trial gives a probability of less than one in a million that the differences are 
due to chance alone.] 


54 


HOVANITZ AND CHANG 


/. Res. Lepid. 


EFFECT OF LARVAL SIZE ON PLANT SELECTION 

Tests for the determination of influence of larval size on plant 
selection were made in the same manner as indicated previously, except 
that all tests were carried out in a room with no wind or breezes. 
Nevertheless, the flat was turned 90° three times for each test. This 
applies to all subsequent tests for the sake of safety. Two . series of 
tests were made, using larvae of different origin. 

The first series of tests (Table 2) was carried out with the use 
of kale-bred larvae. Larvae of different ages and sizes (size is roughly 
proportional to age) were separated and measured. Four larvae were 
isolated, these being the following sizes:: 8.4, 14.7, 19.0 and 21.0 mm 
in length. The tests were carried out by five trials of a given larva 
at each position with respect to the direction of the wind (Fig. 1), 
giving a total of 20 trials for each size larva. 

TABLE 2. Choice of five kinds of plants by different sized larvae 
of Pieris rapae ( Series 1 ) 

Size of Total 

larvae The number of times and percentage larva goes to Number 


(mm) 

Kale 

Mustard 

Radish 

Nasturtium 

Isomeris 

None 

of trials 

21.00 

16 - — 80 % 

4 — 20 % 

0 — 0 

0—0 

0—0 

0—0 

20 

19.00 

13 — 65 % 

3 — 15 % 

3 — 15 % 

1 — 5 % 

0—0 

0—0 

20 

14.70 

11 — 55 % 

4 — 20 % 

4 — 20 % 

1 — 5 % 

0—0 

0—0 

20 

8.40 

10 — 50 % 

5 — 25 % 

5 — 25 % 

0—0 

0—0 

0—0 

20 


One single kale-bred larva was used for each size; tested each larva five times 
at each position. 

The 8.4 mm larva selected kale 50%- of the time, radish 25% of the 
time and mustard 25% of the time. 

The 14.7 mm larva selected kale 55%, mustard 20%,. radish 20% 
and nasturtium 5 % of the time. 

The 19.0 mm larva selected kale 65%, mustard 15%, radish 
1 5 % and nasturtium 5 % of the time. 

The 21.0 mm larva selected kale 80% and mustard 20% of the 
time and no others. 

The probability, as caculated by chi-square, that the differences 
as indicated might be due to chance are less than 1/million, 1/100,000, 
1/500, 1/500 respectively for each experiment. 

CONCLUSIONS: 

( 1 ) The increase in selection of kale for each experiment is 
correlated with increase in size of the larvae ( Fig. 2 ) . 

(2) Kale is the plant selected over all the others in this experi-\ 
ment. It should be noted that the larvae had been bred on kale prior 
to the experiment, and previous generations had been bred on cabbage. 

(3) Not one larva went to Isomeris'. This is interesting in view 
of the observation in the preceding experiments that Isomeris was 


(i); 5 i- 6 i, 1962 


THREE FACTORS 


55 



2. The relation between size (and age) of larvae and the percentage of the 
time kale was selected. The size of the larvae is directly correlated with an 
increase in selection of kale. 


repellent to Pieris rapae larvae. 

(4) The older or larger the larvae, the greater the selection of 
kale over all other plants ( Tig. 2 ) . The relationship is not a straight- 
line, but rather a geometric increase. This indicates that there is an 
increase in the critical powers of food perception in the older or 
larger larvae. 

In the Series No. 2 tests (Table 3 ), larvae were used which had 
previously been grown on nasturtium. They were obtained from 
Ventura County on nasturtium and were continued in the laboratory on 
that plant. 

Two larvae 22.4 mm in length, and four larvae 16.5 mm in length 
were used in this experiment. Otherwise, the trials were managed 
in the same way as in previous experiments with the exception that 
the total number of trials was 80 for large larvae and I 6 O for the 


56 


HOVANITZ AND CHANG 


/. Res. Lepid. 


smaller larvae. 

The selection by these larvae bred on nasturtium (Table 3)' was 
radically different from the selection by larvae bred on kale ( Table 2 ) . 
The selections of kale by the small larvae bred on nasturtium were 
15% as compared with 55% for a comparable size bred on kale. The 
selections of kale by the large larvae bred on nasturtium were 22% 
compared with 80% for a comparable size bred on kale. The greatest 
shift in food plant selections for the nasturtium-bred small larvae 
was 49% to mustard as compared with 25%, 9% to nasturtium as 
compared with 0%, and 16% with no selection as compared with 0%. 
For the large larvae, comparable data were 34% to mustard 
as compared with 20%, 33% to nasturtium as compared 

with 0%, and 9% with no selection as compared with 0%. 
CONCLUSIONS: 

( 1 ) The shifts in food-plant-selection by the larvae grown on 
different plants are highly significant and indicate a direct relationship 
between the feeding habits of the larvae, and their selection when 
given a free choice. 

(2) The direct relationship between the size (and age) of the 
larvae and their selection of the plant previously eaten is clearly 
indicated. 

(3) A plant previously of nearly negligible selection may be the 
most highly selected if the larvae have been grown on that plant. For 
example, larvae selected nasturtium over all others (33%) after being 
bred on that plant (Table 3), whereas larvae bred previously on kale 
had almost no selectivity for that plant. 

INFLUENCE OF PREVIOUS FOOD PLANTS OF LARVAE 

Additional experiments to test the influence of previous exposure 
to larval food plants are reported on here. 

The first comparative test involves two strains of Pieris rapae, one 
from a population growing in the wild on black mustard and the other 
growing on cabbage. These strains were kept in the laboratory for 
several generations; the mustard strain was bred for over six genera- 
tions on mustard in the laboratory and the kale strain was bred on kale 
in the laboratory for over ten generations. 

Larvae were selected for these experiments which had a size of 
14 to 16 mm. Using twenty-five larvae of each strain, 600 tests of 
selection were made (Table 4) : the selected plants were mustard, 
kale, nasturtium, Isomeris and Cleome. 

The only significant differences detected in the tests between the 
two strains were the relative selections made of mustard and kale. 
Larvae from the mustard-bred strain selected mustard 61% of the time 
compared with the kale-bred strains of larvae of 24% on that plant. 
On the other hand, kale was selected by the mustard-bred larvae 20% 


TABLE 3. Choice of fi¥e kinds of plants by different sized larvae 
of Pieris rapae ( Series 2 ) 


57 


1^62 


THREE FACTORS 



. m 



: > 

CM 

d 

! ^ 


Z 



lb, 

, m 


tt 

! .2 

0 0 

jj| 

* ^ 

: © 

» \o 





ffl 

cwo 


0 

1 i 


z 

,u 




0 



0 

01 

m 


© 

m 

E 

0 

II 

m 



© 



ra 

«0 

E 


E 

.2 

mc\ 

© 

u 

""C 

m 


1 

1 1 

■s 

c 

m 

Z 

\o 

,-4 

ffi 



m 

© 

E 





^ 0 

H- 



0 

m 

1 1 

© 



M 



E 



3 



c 

m 

*0 




C\ 

i- 

•S 



m 

1 


1 

1 1 

r- o\ 

CN r^ 






fN| \f\ 



^ T 


£ 

1 

w 

^ CM 

og 


^ fr\ 

m — 

6 

£ 

rvivd 

fNj 1-1 


a 

.S g 

d 

r ’S 

S 3 


« o 


^ s « 

4* y -i 
o « 

z 

S”" o 


OT aj 

^ « 3 

9J 

U 
Ci 
> 


« e 


^ a, S 
« ^ 

t 

-G 

«— « M O 

*u S 3 
^ a . 

S g i 

p o G 

■g 

3 Si5 

d U CN 



^ « flj 

Ms- 

a rt 
o > « 

— I P4 
4> 

•S-a a 
2 3® 
^ ^9 

G ^ m 

""W 2 

1 11 

8-§ « 

ej 'S 

^ bo S 

e 

a m O 


T II 

u ° g s 

8- 

. uO S-l 

^ S p 

S 2 2 S 

^ So «« 

0^0 

OJ ^ a r-l 

2^ S M 

3 «J s > 

^ fe O 
^ ^ 

^ g G '-2 

b 

^ o 

*-**-• 2 y 

P«i 

S 6 8^ 



The nastuf tram-bred larvae were from Ventura County, collected in a nasturtium 
garden. The parents of the kale-bred larvae were collected io western Orange 
County on cabbage; the larva are the first generation fed on kale. 


58 


HOVANITZ AND CHANG 


/. Res. Lepid. 


of. the time as compared with 59% by the kale-bred larvae. Testing 
these differences by chi-square using the four-fold table indicates a 
probability of less than 1 /million that the differences could be due to 
chance alone. This may be calculated in another way, that is, by taking 
364 + 354 = 718 for one value and 119 + 144 = 263 for the other. 
The expectation that these differences might be due to chance alone, 
that is, that the deviations 718 — 981, and 971 — 263 are not due to 

2 2 ‘ 
chance is well over one in a million. 

CONCLUSIONS: 

( 1 ) The shift in selection of mustard or of kale according to 
whether or not the larvae were from a kale or mustard strain is 
indicated by the experiments. The data are highly significant at an 
incredibly high level. 

( 2 ) There is no significant change indicated in the selective level 
of the other plants involved as to whether or not the larvae were from 
the mustard or kale strain. 

The second series of data on the problem of food plant selections 
involves nasturtium-bred larvae from a nasturtium strain and kale- 
bred larvae from a cabbage strain (Table 5). The experiments were 
carried out as before, two larvae of each strain were used, the same 
size for a total of forty trials each, of which ten were in each position 
as indicated in Figure 1. 

The differences between the plant selections are again highly 
significant. The nasturtium -bred larvae as compared with the kale- 
bred larvae preferred mustard (35% as compared with 22.5%) and 
nasturtium (15% over 4%). The kale-bred larvae preferred kale 
(49% over 12.5%) and radish (22.5% over 14.0%). Neither 
selected Isomeris to any extent. A high proportion (22%) of the 
nasturtium-bred larvae rejected all plants. 

CONCLUSIONS: 

( 1 ) As in previous experiments, there was a shift in selection by 
the larvae according to the previous larval food plants. 

( 2 ) This shift in selection was in the direction of the plant they 
had previously fed on. For example, nasturtium-bred larvae selected 
nasturtium better than did kale-bred larvae. 

( 3 ) Even though nasturtium-bred larvae selected nasturtium 
better than did kale-bred larvae, mustard was preferred over nasturtium 
and over kale. 

(4) The greater selection of mustard over kale bv nasturtium- 
bred larvae is significant not only in this experiment but also in a 
previously indicated test (Table 3). In both cases, kale is poorly 
preferred as compared with mustard when the larvae were nasturtium- 
bred. 

(5) Radish is of lower selective influence when the larvae are 


(i): 5 i- 6 i, 1962 


THREE FACTORS 


59 


nasturtium-bred than when they are kale-bred. This was indicated 
also in previous tests ( Tables 3 and 1 ) . 

(6) A significantly higher percentage of rejects (22%) are 
made by the nasturtium-bred larvae than by the kale-bred. This was 
also indicated in previous tests (Table 3 as compared with 1, 2 or 4). 

DISCUSSION 

All three factors, investigated as to their relevance in influencing 
larval choice of food, have been shown to be important in that regard. 
These factors are: 

( 1 ) Influence of air movemenr. 

(2) Influence of age (size) of larvae. 

(3) Influence of food previously eaten by larvae, or strain of 
larvae. 

In addition, by virtue of the laboratory set-up used in these 
experiments, some further conclusions may be drawn which are not 
completely in accord with some previous conclusions reached by others. 
One of these further conclusions is with regard to the ability of 
lepidopterous larvae to "recognize” food plants from a distance. In the 
present experiments, the larvae were attracted to the plants from a 
distance of over 120 mm without any difficulty and responded in most 
cases immediately. 

Chin (1950) reports that the larvae of the Colorado potato beetle 
(Leptinotofsa) could not perceive its food plant from a distance farther 
than 2mm. Gupta and Thorsteinson (I960) tested the olfactory 
response of the larvae of the diamond backed moth [Plutella macul- 
ipennis (Curt.)] toward mustard oil odors and found that 5 mm was 
the maximum distance of response. Dethier (1959) in a study on 
food plant distribution and density and larval dispersal as factors 
affecting insect populations, states in relation to field observations that 
"Larvae find new food-plants by chance alone.” It is presumed that 
he had reference to the gross search for new food plants, not the 
closer relationship. In the case of the experiments of Chin, and Gupta 
and Thorsteinson, the screen test was used. In this, the larvae were 
placed on a screen above the food plant or other attractive source. In 
the present experiments, the larvae were below the food plants. This 
difference could account for a fundamental difference when testing for 
an odiferous substance heavier than air. Under natural conditions, our 
observations have always indicated that larvae tend to climb upwards 
rather than down so that tests involving simulated natural conditions 
should be best performed with the attraction above rather than below 
the larvae (Fig. 3). This point will be covered more fully in tests 
to be described in later papers. 

In a paper available to us after the tests here indicated were 
performed, (Takata, 1959), there has been indicated a change in food 
plant preference of larva when reared successively on different food 


60 


HOVANITZ AND CHANG 


/. Res. Lepid. 



3. Larva of Pieris rapae on edge of pot during one of the selection experiments 
reaching out to black mustard plant three to four inches away. The larva 
had already traveled several inches under directed impulse to get this far. 


plants for several generations. These results complement ours, even 
though the technique of testing used by Takata was different. He used 
cabbage and radish with Pieris rapae crucivora larvae. In testing, he 
used actual feeding experiments of the weight of leaves of the plants 
eaten by the larvae. 


(i): 5 i- 6 i, 1962 


THREE FACTORS 


61 


SUMMARY 

The evidence in this paper indicates ( 1 ) that larvae of Pieris rapae 
are capable’ of detecting their preferred food plant from a distance of 
at least 120 mm under the conditions of these experiments, (2) that 
the direction of wind movement is very important to detection of food 
plant, presumably through odor, (3) that the older and larger larvae 
are much better at selecting the preferred food plant at least at a 
distance and (4) that the influence of food previously eaten by the 
larvae, or the strain of the larvae, directly influences their choice of 
food in the direction of the previously eaten food. 


LITERATURE CITED 

CHIN, CHUN-TEH. 1950. Studies on the physiological relations between 
the larvae of Leptinotarsa decemlineata Say and some solanaceous plants. 
H. Veenman and Sons, Wageningen, Holland, pp. 1-88. 

DETHIER, V. G. 1959. Food plant distribution and density and larval 
dispersal as factors affecting insect populations. Canad. Entom. 91: 581-596. 

GUPTA, P. D. AND A. J. THORSTEINSON. I960. Food plant relation- 
ships of the diamond-backed moth (Plutella maculipennis (Curt.)). I. 
Gustation and olfaction in relation to botanical specificity of the larva. 
Ent. exp. and appl. 3 (I960) :24l-250. 

TAKATA, N. 1959. Studies on the hobt preference of common cabbage 
butterfly, Pieris rapae crucivora (Boisduval). VI. Change in the food 
preference of larvae when reared successively by the definite food plant 
for several generations (preliminary report). Jap. J. Ecol. 9:224-227. 



1 ( 1 ) : 63 - 71 , 1962 


Journal of Research on the Lepidoptera 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
Copyright 1962 


THE GENERIC, SPECIFIC AND LOWER CATEGORY 
NAMES OF THE NEARCTIC BUTTERFLIES 

PART 1 — The Genus Pieris 

PADDY McHENRY 

1032 E. Santa Anita, Burbank, California 

PREFACE 

INTRODUCTION 

The author has asked that I write a brief introduction to this, 
the first of a series of compilations of generic, species and lower 
category names of the Nearctic butterflies. The objects of the work are 
(1) to give a complete list of all the names proposed for the butterflies of 
the Nearctic region from the year 1738 until the date of publication 
of the list, (2) to indicate the date and place where originally published, 
(3) to indicate the types of the genera, species and lower category 
names where known, (4) to indicate the type locality in the original 
author’s words for types, except those for the generic names, and 
(3) to indicate the location where types have been deposited where 
indicated by the author. 

The biological application of these names so indicated is not to 
be attempted in this series. Instead, this effort is left to the individual 
who makes a comprehensive study of the biology of the groups con- 
cerned, such a person rarely having the time and inclination to make 
the detailed and laborious studies of nomenclature as such. There 
is urgent need for a work such as this even in a field as well civilized 
as the Lepidoptera; in fact, this kind of work is so long overdo that 
the civilized state is rapidly reverting to chaos. Each new volume 
which appears from the press on this group of insects plows under well 
established names with no reason given to justify such destruction. 
A most valuable start in the direction of making reason out of chaos 
was started by Francis Hemming in his TThe Generic Names of the 
Holarctic Butterflies” published in 1934. This work is proceeding 
also in the sam.e direction. 

When the various parts of this work are completed, it is expected 
to reissue the whole as a separate volume. Then, perhaps we can 
prevail upon Mr. McHenry to go on to a similar effort with the moths, 

William Hovanitz 


63 


64 


McHENRY 


/. Res. Leptd. 


Each entry for the generic names includes the following; the 
name, the author’s name, the most accurate determinable publication 
date, the bibliographical reference (including the volume or subvolume 
for the date indicated ) , the type species, the selection of the type species 
and the supporting data to justify the type indicated. 

Each entry for the specific names includes the following; the name 
as originally given, the author’s name, the most accurate determinable 
publication date, the bibliographical reference, data on type locality, sex, 
season, etc., other spellings of the name and notations of any homonym. 

The date of publication of a name is indicated in a manner that 
shows its relative accuracy; for example, "mailing” and "release” dates 
if known are given preference over "publication” dates if they are not 
the same. All dates are given in English and normally in the sequence 
of day, month and year. Where more than one date is indicated, the 
more accurate one is given after the less accurate one. Dates ascertained 
directly from a source whose purpose was the determination of time of 
publication of a work are given without brackets; others, ascertained less 
directly are given in brackets and explained in a footnote. Combinations 
of dates with and without brackets may be given when necessary. 

I have seen originals or photostats of all references except as noted 
but have been unable to ascertain the contents and dates for the various 
parts of Verity’s Rhopalocera Palaearctica. 

The names of the genus Pieris are listed under the five species 
indicated, since in this genus the separation does not involve great 
difficulty. I have been aided in this regard by William Hovanitz, or 
by reference to the original description. Should these be in error, 
no harm is done since the total list, not the biological interpretation of 
the categories to which the names applied, is the purpose of the listing. 

Should anyone find names which have been inadvertently omitted 
from this listing, a great service will be rendered to so notify the author 
or the editor of this fact so that corrections may be made. 

I take this opportunity to express my gratitude for the use of 
facilities and to the staffs at the Allan Hancock Foundation Library of 
the University of Southern California, and at the Los Angeles County 
Museum Library & Entomological Department. A debt of gratitude is 
tenderd to the photographic service sections of the U. S. Department of 
Agriculture Library, Harvard College Library, New York Public Library, 
University of California Library and Cornell University Library. An 
extended exchange of correspondence with Mr. C. F. DosPassos has 
been most helpful. 

LIST OF GENERIC NAMES USED OR 
AVAILABLE FOR PIERIS 

ASCIA Scopoli. 

Type; monuste L. 


(i):63-7I, 19^2 


NAMES Pt. 1 


65 


MANCIPIUM Hubner. 

Type: hellica Hubner = hellica L. = helice L. 

PIERIS Schrank. 

Type: hrassicae of var. authors = brassicae L. 

ANDROPODUM Hubner. 

Type: brassicae Hubner = brassicae L. 

GANORIS Dalman. 

Type: brassicae L. 

TACHYPTERA Berge 

Type: brassicae L. 

PONTIA Fabricius. 

Type: daplidice Fab. = daplidice L. 

SYNCHLOE Hubner. 

Type: callidice Butler = callidice Esper = callidice Hubner 

ASCIA SCOPOLI. 1777. Introd. Hist. Natur. : 434, no. 175. Includes 
among others no. 80 (="P. D. [C.] monuste” of Linnaeus, 1767) 
which in turn is Papilio D. C. monuste Linnaeus, 1764. 

Type Papilio D\anaus\ C[andidus]. monuste Linnaeus. 1764. Museum 
Ludovicae Ulricae ([I]) : 237, no. 56. 

Type Selection. Scudder. 1872 [in or pre-June] L Syst. Rev. Some Amer. 
Butt., p. 40. Says of Ascia: "Type Papilio monuste Linn.” 

MANCIPIUM HUBNER. [1 Jan. 1807] - [19 Dec. 1807]L Samrn. Exot. 

Schmett, 1: pi. [141], ff. 1-4. Gives "Mancipium vorax hellica”. As 
used by Hubner, hellica is considered to be hellica Linnaeus, 1767 
which in turn is helice Linnaeus, 1764. 

Type. Papilio D[anaus]. C[andidus]. helice Linnaeus. 1764. Museum 
Ludovicae Ulricae ([I]) : p. 243, no. 62. 

Type Selection. Hubner. As above. Gives only hellica, at the time, 
which becomes the type. 

PIERIS SCHRANK. 1801. Fauna Boica 2(1): 152, no. 198; 2(1): 

160-170, nos. 1283-1296. Includes Pieris brassicae which he says is 
Papilio brassicae of Goeze, Borkhausen, Brahm, and Esper. Of these 
authors, brassicae is considered to be hrassicae Linnaeus, 1758. 

Type. P\ap)ilio\ D\anaus\ [Candidas] hrasicae Linnaeus. 1758. Syst. 

Nat. 10th Ed. 467-468, no. 58. 

Type Selection. Latreille. 1810. Consid. Gen. Anim. Crust. Arach. Ins. 
p. 440. Says: "Pieride. Pontia brassicae, Pab.”. As used by Latreille, 
brassicae Fabricius !s considered to be brassicae Linnaeus, 1758. 

ANDROPODUM HUBNER. 1822. [post 22 Sept.]." Syst.-Alph. Verz. pp. 

2-5, 7-9. Includes "Brassicae L, 401-403. Andropodum vorax.” 

Type. P\apilio\ D[anaus]. [Candidas] hrassicae Linnaeus. 1758. Syst. 
Nat. 10th. Ed. 1:467-468, no. 58. 

Type Selection. Hemming. Sept. 1933. Entomologist: 66(844) : 199. 
Says of Andropodum: "Type =: Andropodum hrassicae Linn.” 

GANORIS DALMAN. 1816. Kongl. Vetenskaps Academiens Handlingar 
1816(1) : 61-62, no. 8; 1816(1) : 86-90, nos. 1-10; tablet 1, no. 8; 
tablet 2, no. 8. Includes G. brassicae in which he cites Linnaeus 
Fauna Sv., pp. [269] -[2701, no. [10351 (this is hrassicae Linnaeus, 
1761, which is hrassicae Linnaeus, 1758). 

Type. Plapilio]. D[anaus]. [Candidas] Brassicae Linnaeus. 1758. Syst. 
Nat. 10th Ed. 1:467-468, no. 58. 

Type Selection. Dalman. As above. Says of Ganoris: "Generis Typus: Pap- 
hrassicae”. 


66 


McHENRY 


/. Res. Lepid. 


TACHYPTERA BERGE. 1842. Schmetteclingsbuch, p. 19, pp. 92-105. 
Not seen, given as per Hemming, 1934. 

Type. P[apilid\, .Dlanaus\ [Candidus] brassicae Linnaeus. 1758. Syst. 
Nat. 10th. Ed. 1:467-468, no. 58. 

Type Selection. Hemming. Feb. 1934. Entomologist: 67(849) :38. Says 
of Tachyptera: "Type Papilio brassicae Linn., 1758.” 

PONTIA FABRICIUS. 1807. In Illiger. Magazin fur Insektenkunde. 6: 

283, no. 23. Includes: ’Tap . , . Daplidice”. As used by Fabricius 
daplidice is considered to be daplidice Linnaeus, 1758. 

Type. P[apilio]. D[anaus]. [Candidus] daplidice Linnaeus. 1758. Syst. 
Nat. 10th Ed. 1:468, No. 2. 

Type Selection. Curtis. 1824. Brit. Entom. 1: pi. 48. Not seen, 
given as per Hemming, 1934. 

SYNCHLOE HUBNER. 1818 [post 22 Dec.]^. Zutr. Samm. Exot. Schmett. 

1st. 100 (text in 1): p. 26, under no. 76. (Synchloe autodice). 
Includes "S. callidice”. Refers to his earlier publication of this 
butterfly. 

Type. Papilio callidice Hifbner. [24 Dec. 1799] - [13 Apr. 1800]^. Samm. 
Europ. Schmett. (Papiliones) : plate 81, figs. 408-409. Page 63, no. 5 
and plate 108, figs. 551-552 were published later. 

Type Selection. Butler. 12 Sept. 1870. Cistula Ent. 1(3):51. Says of 
Synchloe: "Type S. callidice Esper”. As used by Butler callidice Esper 
is considered to be callidice Hubner [24 Dec. 1799] - [13 Apr. 1800]. 

LIST OF SPECIES AND LOWER CATEGORY NAMES 
USED OR AVAILABLE FOR PIERIS 

1. P. (PONTIA) BECKERII EDWARDS. 
heckerii Edwards. 

gzmderi Ingham. 
pseudochloridice McDimnough. 

2. P. (PONTIA) SISYMBRII BOISDUVAL. 
elivata ( B. & B. ) . 

jlava Edwards. 
flavitincta Comstock, J. A. 
sisymhrii Boisduval. 
transversa {B. 8iP>.) 

3. P. (PONTIA) PROTODICE BOISDUVAL & LECONTE 
calyce Edwards. . 

nastuftii Edwards ( Bdv. Ms. ) . 
nehoni Edwards. 
occidentalis Reakirt. 
protodice Boisduval & LeConte. 
vernalis Edwards. 

4. P. (PIERIS) NAPI (LINNAEUS). 
acadica Edwards. 

aestiva Edwards. 
arctica Verity. 
borealis Grote. 
casta Kirby. 


i(i):6}-7T, 1962 


NAMES — Pt. 1 


67 


castoria Reakirt. 
cottlei Gunder. 
cruciferamm Boisduval. 
flava Edwards. 
flava Edwards. 
ffigida Scudder. 
hulda Edwards. 
hyemalis Edwards. 
iberides Boisduval. 
macdunnoughii Remington. 
marginalis Scudder. 
micro striata Comstock, J. A. 
mogollon Burdick. 
napi (Linnaeus). 
nasturtii Boisduval. 
ochsnheimeri Staudinger. 
oleracea Harris. 
pallida Scudder. 
pallidissima B. & McD. 
pseudobryoniae Verity. 
pseudoleracea Verity. 
pseudonapi B. & McD. 
resedae Boisduval. 
venosa Scudder. 
virginiensis Edwards. 1870. 
virginiensis Edwards. 1881. 

5. P. (PIERIS) RAPAE (LINNAEUS). 
aestivus Verity. 

immaculata De Selys-Longchamps. 
immaculata Cockerell. 
immaculata Skinner & Aaron. 
metra Stephens. 
novangliae Scudder. 
rapae ( Linnaeus ) . 
yreka Reakirt. 


1. PIERIS (PONTIA) BECKERII EDWARDS. 

heckerii Pieris Edwards. [Sept. 1871]*^. Butt. N. Amer. 1(8) : [28a] - 

[29]; plate [81], figs. 4-7. 'cf described. "Virginia City, Nevada, 

April 1870.” "Four individuals”. McDunnough, 1 Dec. 1928, spells 
heckeri. 

gunderi, Ascia heckerii Ingham. 14 Apr. 1933. Pan-Pacific Ent. 9(2) :75. 
Holotype ? : "Bouquet Canyon, Los Angeles County, Calif”. "Bred 
from larva”, "emergence date June 21, 1932”. Type deposited: Calif. 
Acad. Sci., San Francisco, Calif. 

pseudochloridice, Pieris heckeri McDunnough. 1 Dec. 1928. Can. Ent. 
60(11) : 266-267. Holotype d* : "Oliver, B[ritish]. C[olumbia]., April 24 
. . . No. 2861 in the Can. Nat. Coll., Ottawa”. Allotype ? : "Oliver, 


68 


McHENRY 


/. Res. Lepid. 


B.C., April 22 ”. Paratypes — I d*: "Hedley, B. C., May 15”. 1 9 : 
"Osoyoos, B.C.” 1 + : "Osoyoos, B.C.” 

2. PIERIS (PONTIA) SISYMBRII BOISDUVAL. 

elivata, Ascia sisymhni Barnes & Benjamin. 8 Dec. 1926. Bull. Sou. Calif. 
Acad. Sci. 25(3) :88, no. 33a. "Type locality: Glenwood 
Springs, Colo.”. "Number and sexes of types: Holotype d* May 1895, 
Allotype ? May 1895, 7 d» 14 9 Paratypes, various dates April to 
June”. 

jlava, Pieris sisymhri Edwards. Apr. 1883. Butt. N. Amer. 2(11): [67]; 

plate [151. fig 5. ? described. No locality, series data nor dates given. 
flavitincta, Pieris sisymhrii Comstock, J. A. 20 Feb. 1924. Bull. Sou. Calif. 
Acad. Sci. 23(1) : 19-20; plate 7, fig. 9. 9 described. "The example 
before us was captured . . . April 30th, 1911 at Cranbrook, British 
Columbia”. Intends to propose new name for jlava Edwards. 
sisymhrii, Pieris Boisduval. 1852 [25 Feb. - 22 Dec. Ann. Soc. Ent. 
France. 2nd Ser. 10 (2) :284, no. 8. 9 described. [California]. No series 
data nor dates given. Edwards, Apr. 1883, spells sisymhri. 
transversa, Ascia sisymhrii Barnes & Benjamin. 8 Dec. 1926. Bull. Sou. 
Calif. Acad. Sci. 25(3): 88, under no. 33. "Type localities: Paradise, 
Cochise Co., Ariz.; Redington, Ariz.”. "Number and sexes of types: 
Holotype cf , Allotype 9,1 9 Paratype, all March; 1 cf Paratype, no 
date”. 

3 . PIERIS (PONTIA) PROTODICE BOISDUVAL & LECONTE. 

calyce, Pieris Edwards. Nov. 1870. Trans. Amer. Ent. Soc. 3(?) sign. 25: 
189, no. 1 (Nomen nudum); 189. d* described. "From Nevada; in the 
collection of Henry Edwards”. No series data nor dates given. 
nasturtii, Pieris Edwards (Boisduval Ms. name). [9 May 1864]°. Proc. Ent. 
Soc. Phila. 2(4) : 501, no. 1 (Nomen nudum); 501. d* & 9 described. 
"San Francisco; from Dr. Behr, who informs me that it is common in 
some localities in the vicinity of that city”. No series data nor dates 
given. 

nelsoni, Pieris Edwards. Apr. 1883 Butt. N. Amer. 2(11): [71] - [72]; 

plate [15], figs. 6-7. "1 cf ... St. Michael’s Alaska, June, 1881”. 
occidentalis, Pieris Reakirt. June 1866. Proc. Ent. Soc. Phila. 6 (pp. 
121-152) : 133-134. cf & 9 described. "Hab. - Rocky Mountains, 
Colorado Territory, California, (Coll. Tryon Reakirt)”. No series data 
nor dates given. 

protodice, P[ieris]. Boisduval & LeConte. 1829. Hist. Gen. Icon. Lepid- 
Chen. I’Amer. Sept. 1 (5) :45-46. Plate 17, figs. 1-3 published later. A 
& 9 described. "Elle parait au printemps et a la fin de juin aux 

environs de New-York. Nous en avons vu un individu pris dans le 

Connecticut”. 

vernalis. Pieris Edwards. [9 May 1864]®. Proc. Ent. Soc. Phila. 2(4) : 501, 
no. 2 (Nomen nudum); 2 (4) :501-502. cf & 9 described. "In the 
collection of Mr. Geo. Newman and Mr. Wilt are several specimens, 
taken, as I am informed, at Red Bank, New Jersey, in the month of 
May”. "There is also in the Society’s collection a pair from the Rocky 
Mountains, that appear to be identical with it”. 

4. PIERIS (PIERIS) NAPI (LINNAEUS). _ . 

acadica, Pieris napi Edwards. June 1881. Papilio l(6):86-88 (in pt.), 

sub no. 1; p. 98, no. 1 of no. 3; p. 99; plate 3, figs. 10-11. It appears 
that one pair was from Southern Newfoundland and was taken in the 

last week in July, it also appears another set ( 1 cf 2 9 ) had emerged 

from pupae in August and was from the same locality. 
aestiva, Pieris napi oleracea Edwards. June 1881. Papilio l(6):86-88 (in 
pt.. Nomen nudum), sub no. 1; l(6):89-95 (in pt.) 

sub no. 3; l(6):95-98 (in pt.), sub no. 4; 1(6):98, no. 3 of no. 3; 


i( 1962 


NAMES — Pt. 1 


69 


1(6):99; plate 3, figs. 15-16. cf & described. "NewYork” (see 
figs, explanation, p, 99). "summer brood”. No series data given. 
arcUca, Pieris napi frigida Verity. 1905-1911. Rhopal. Palaearctica : pp. 
333-334 (in pt., cites figs 32-33 on plate XXXII and figs. 16-17 on 
plate LXVII as P. n. f. arctica)\ plate XXXII, figs. 32-33; plate 
XXXII explanation page, figs. 32-33 (gives as P. n. frigida); plate 
LXVII, figs. 16-17; plate LXVII explanation page, figs. 16-17 (gives as 
P. n. arctica); p. xxviii (cites figs. 31-33 & 36-37 on plate XXXII 
and figs. 16-17 on plate LXVII, gives as P. n. /. arctica). "31 . . . cf 
(ile d’Yesso, Japon)”. "32 . . . c? , (Norvege sept.)”. "33 . . . d* 
(Nulato, Alaska)”. "36 ... $ (Nulato, Alaska)”. "37 ... 9 
.(Finmark, Scandinavie) ”. ”16 , 9 (Saltdalen, Norvege sept.) ”. "17 

. . . ? (Laponie)”. Figs. 31, 33, 36: [coll, de Joannis]. Fig. 32: [coll. 
Stefanelli]. Fig. 37: [coll. Obth.] Fig. 16: le coll. Mourray]. Fig. 17: [e 
coll. Leach]. No series data nor dates given. 
borealis, Ganoris oleracea Grote. Nov. 1873. Bull. Buffalo Soc. Nat. Sci. 
1(4) sign. 24:185. "months of June and July, on the Island of Anti- 
costi”. No series data nor sex given. 

casta, Pontia Kirby, W. 1837. In J. Richardson. Fauna Boreali-America 
(4) :288, no. 1; plate 3, fig. 1. "Three specimens taken in Lat. 65°”. 
No sex given. 

castoria, Pieris Reakirt. [11 June 1866] - [13 Feb. 1867]*^. Proc. Acad. Nat. 
Sci. Phila. [18] (3) :238, no. 2. d* described. "Hab. - California. Coll. 
Tryon Reakirt”. No series data nor dates given. 
cottlei, Pieris napi castoria Gunder. 6 July 1925. Ent. ^ews 36(7) :197- 
198, no. 9; plate 5, fig. 9; Holotype d : "(author’s Coll.), Anderson 
Springs, Lake County, California, May 5, 1919”. 
crucifer arum, Pieris Boisduval. 1836 [in or post Apr.]^°. Hist. Nat. Ins. 
Spec. Gen Lepid. 1 (text in 1) :519, no. 119. d* described, "dans les 
provinces septentrionale des Etats-Unis”. No series data nor dates given. 
flava, Pieris napi pallida Edwards. June 1881. Papilio 1(6) :89”95 (in pt., 
unnamed), sub no. 3; 98, sub no, 2 of no. 3. "Washington Terr[itory].”. 
"One of these [a $ ] is yellow on the upper side”. No date given. 
flava, Pieris napi venosa Edwards. June 1881. Papilio 1(6) :88-89 (in pt,, 
unnamed), sub no. 2; 98, sub no. 1 of no. 2. Fig. 7 of plate 2 
(although not so named in plate explanation, p. 99) appears to be 
flava. "A large percentage of female Venosa are yellow on the upper 
side”. No locality, series data nor dates given. 
frigida, Pieris Scudder. Sept. 1861. Proc. Boston Soc. Nat. Hist. (8(?) 
sign. 12:181-182. "Two . . . males . . . two females”, "on Caribou 
Island, Straits of Belle Isle”. No dates given. 
hulda, Pieris Edwards. Sept. 1869. Trans. Amer. Ent. Soc. 2(?) sign. 

48:370. "From Kodiak, 1 o* . Coll, of Henry Edwards”, No date given. 
hyemalis, Pieris napi oleracea Edwards. June 1881. Papilio l(6):88-89 
(in pt.. Nomen nudum), sub no. 2; 89-95 (in pt.), sub no. 3; 95-98 
(in pt.), sub no. 4; 99; plate 2, fig. 8. d* &■ $ described. Directly and 
indirealy cites material from New Hampshire, Massachusetts, Lake 
Superior area and Mt. Hood, Oregon. No series data nor dates given. 
iherides, Pieris Boisduval. [1869, pre 1 Nov.]^’. Ann. Soc. Ent. Belgique 12 
(in 1):39, no. 9. d* & ? described. [California]. No series data nor 
dates given. 

macdunnoughii, Pieris napi Remington. 17 Sept. 1954. The Lepid News 
8(3-4) :75. A new name for Pieris napi pseudonapi Barnes & McDun- 
nough, 5 Dec. 1916. See pseudonapi for series data. 
marginalis, Pieris Scudder. Stpt, I 86 I. Proc. Boston Soc. Nat. Hist. 8(?) 
sign. 12:183. "(Id* ,1 9 ) which are in the Museum of Comparative 
Zoology”. The male . . . from Gulf of Georgia, and the female from 
Crescent City, California”. No dates given. 


70 


McHenry 


/. Re$. Lepid. 


mkrostfiata, Pieris napi Comstock, J. A. 12 Sept. 1924. Bull. Sou. Calif. 
Acad. Sci. 23(4) :125. Holotype f:- "Eldredge, Sonoma County, Cali- 
fornia. March 13, 1911”. Allotype [•$■]: “same locality and on the 
same date”. 'Taratype; one male - same locality and date”. 

mogollon, Pieris napi Burdick. 28 Sept. 1942. Can, Ent. 74 (8) :154-155. 
Holotype : "Mogollon Range, Catron Co., New Mexico, 5»7-40. No. 
5222 in the Can . . . Nat . , . Coll . . . , Ottawa”. Allotype $ : "same 
data”. "Paratypes. 2 ^ ,same data; 3 cf , Sierra Blanca, Lincoln Co., 
N. M., 5-25-40; all in the Can , . . Nat . . . Coll . . . , Ottawa. 1 d* , 
Mogollon Range, Catron Co., N. M., 5-7-40 and 1 $ , Sierra Blanca 
Range, Lincoln Co., N. M., 5-20-40; in the U , . . S . . .^at . . . Mus . . 
Fifty-six paratypes from the above locations in the collection of the 
author, from which other museums will be supplied”. 

napi, P[apilio]. Dlanausl. [Candidus] Linnaeus. 1758. Syst. Nat. 10th. Ed. 
1 ;468, no, 60. No locality, series data, sex nor dates given. 
Edwards, June 1881, spells napae. 

nasturtii, Pieris Boisduval. [1869, pre 1 Nov.]^L Ann. Soc. Ent. Belgique 
(12) (in 1) :38-39, no. 7. described, "dans les champs decouverts au 
pied de la Juba”. "Nous o’avons vu que des males”. No series data 
dates given. A homonym of Pieris nasturtii Edwards, [9 May 1864]. 

ochsenheimeri, Pieris Staudioger. End Apr. 1886. Ent. Zeitung Stettin 
47 (4-6) : 199-200. d & 9 described, "wo sie die beiden Haberhauer in 
Anzahl Ende Juni bei Namagao, jedenfalls hoch in den Gebirgen ge 
fangen batten. Auch Maurer sandte mir in demselben Jahre ein im 
Alai-Gebirge (sudlich von Margelan) gefangeoes + ein”. No series 
data given. 

oleracea, Pontia Har^fs. 10 July 1829. The New England Farmer & Horti- 
cultural Jour. 7(51) :402. "Habitat ... in New Hampshire, and Massa- 
chusetts”. No series data nor sex given, "two broods in a season”: 
[May- June - August]. 

pallida, Pieris Scudder. Sept. 1861. Proc. Boston Soc. Nat. Hist. 8 (?) sign. 
12:183. "five specimens (3 d* » 2 9.), which are in the Museum of 
Comparative Zoology”. "Gulf of Georgia”. No dates given. 

pallidissima, Pieris napi Barnes & McDunnough. 5 Dec. 1916. Contrib. 
Nat. Hist. Lepid N. Amer. 3(2) :59; 142; plate 6, figs. 4-5 & 10. 
"second generation (July, August)”, "type cf and $ from Provo, 
Utah”. No series data given. 

pseudobryoniae, Pieris napi frigida Verity. 1905-1911. Rhopal.- Palaearctica : 
p. 146 (cites figs. 46-37 on plate XXXII, gives as P. n. f. pseudo- 
bryoniae); plate XXXII, figs. 36-37; plate XXXII explanation page, 
figs. 36-37 (gives as P. n. frigida); p. xxviii (cites only fig. 37, gives 
as P. n. f. arctica pseudobryoniae) . "36 ... 9 (Nulato, Alaska) [coll, 
de Joannis]”. "37 . . (Finmark, Scandiriavie) [coll. Obth.]”. No 
series data nor dates given. 

pseudoleracea, Pieris napi frigida Verity. 1905-1911. Rhopal. Palaearaica: 
p. 146; plate XXII, fig. 38; plate XXXII explanation page, fig. 38; 
p. xxviii. "38 . . . d' (Labrador) [e- coll. Boisduval in coll, obth.]”. 
No series data nor dates given. 

pseudonapi, Pieris napi Barnes & McDunnough. 5 Dec. 1916. Contrib. 
Nat. Hist. Lepid. N. Amer. 3(2) :57; 142; plate 6, figs. 1-3. "type ^ 
and 9 taken at Silverton, Colo ... in the last week of July”. No series 
data given. A. homonym of Pieris melete pseudonapi Verity, 1905-1911. 

resedae, Pieris Boisduval. [1869, pre 1 Nov.]^ ’ Ann. Soc. Ent. Belgique 
12 (in 1) :38-39, no. 8. 9 described. "Nous ne connaissons cette espece 
que par un seul individu 'femelle pris . . . sur les bords du Sacramento”. 
No date given. 

venosa, Pieris Scudder. Sept. 1861. Proc. Boston Soc. Nat. Hist. 8(?) 


i(i):6}-ji, 1^62 


NAMES — Pt. 1 


71 


sign. 12:182-183. "twenty specimens (5 cf , 15 9 ), brought to the 
Museum of Comparative Zoology, from San Mateo and Mendocino city, 
California”. No dates given. 

vkginiensis, Pieris Edwards. Jan. 1870. Trans. Amer. Ent. Soc. 3(?) sign. 
2:10, no. 5 (Nomen nudum); 13-14. cf & $ described. "Not uncom- 
mon in the Kanawha district [West Virginia] in the month of May, and 
there replacing Oleracea. I have received from Mr. Saunders occasionally 
specimens taken by him at London, Canada”. No series data given. 
virginiensis, Pieris napi oleracea hyemalis Edwards. June 1881. Papilio 
1 (6) :95-98 (in pt.), sub no. 4; 98, under no. 2 of no. 2. " . . . Virgin- 
iensis . . . has become a true species, although unquestionably, in a higher 
latitude, it appears as an occasional aberration only of Oleracea". Speci- 
men data is obscure. A homonym of virginiensis Edwards, Jan. 1870. 

5. PIERIS (PIERIS) RAPAE (LINNAEUS). 

aestivus, Pieris rapae Verity. 16 May 1913. Jour. Linn. Soc. (London), 
Zool. 32 (215) :177-178. "summer brood”. No locality nor series data 
given. 

immaculatag Pieris rapae De Selys-Longchamps. 1857. Ann. Soc. Ent. 
Belgique 1 (in 1) :5, under no. 5. [Belgique]. No series data, sex nor 
dates given. 

immaculata, Pieris rapae Cockerell. Apr. 1889. Entomologist: 22(311) :99. 
No locality, series data, sex nor dates given. Cites p. I6l in Newman’s 
Brit. Butt. A homonym of Pieris napi immaculata De Selys-Longchamps, 
1857. 

immaculata, Pieris rapae Skinner & Aaron. 2 July 1889. Can. Ent. 21(7) : 
128-129. "five specimens in the collection of Am. Ent. Soc., Dr. Skin- 
ner and E. M. Aaron”. No sex nor dates given. A homonym of Pieris 
rapae immaculata De Selys-Longchamps, 1857 and Cockerell, Apr. 1889. 
metra, Pontia Stephens. 1 Aug. 1827. Illust. Brit. Ent. 1(?) sign, d: 19-20, 
no. 4:146-147 published later, cf & ? described. "The insect occurs 
early in April and a second time towards the end of June. I obtained 
specimens of the first brood at Hertford; and of the second I captured 
some this season, at Ripley at the latter period”. 
novangliae, Ganoris rapae Scudder. Apr. 1872. Can. Ent. 4(4) :79. "Two 
of my specimens were taken in June and July, 1869 — - one ... in 
Shelbourne, N[ew]. H[ampshire] . . . the other ... in Waterville, 
M[aine]., and all are males”. 

rapae, P[apilio]. D\anaus\ [Candidusi Linnaeus. 1758. Syst. Nat. 10th. Ed. 

l(in 1):468, no. 59. No locality, series data, sex nor dates given, 
yreka, Pieris Reakirt. [11 June 1866] - [13 Feb. 1867]'^. Proc. Acad. Nat. 
Sci. Phila. [18](3):238, no. 1. cf & 9 described. "Hab. - California. 
Coll. Tryon Reakirt”. No series data nor dates given. 

lAmer. Nat. 6: 3S4-559. Work is reviewed in June 1872 number. 

^Hemming, 1937. Hubner 1:327-437. 

^The work title page signature date qualified by preface signature date on page vi. 

•^Title page signature date qualified by preface signature on page 6. 

■^Hemming. 1937. Hubner. 1:146-324. 

^Hemming. 1931. Proc. Ent. Soc. London. Ser. A. 6: 42-45. Gives actual dates of publication for 
the parts of Edwards’s Butt. N. Amer., (Ser. I). 

7Tome 10 title page signature date qualified by the meeting date of 25 Feb. (p. 275) and the 
evidence indicating that the tome to page 489 was published before the meeting of 22 Dec. 
(p. Ixxxii in Bulletin). 

SProc. Ent. Soc. Phila. 1864. 3: p. 695. Notes that No. 4 of Vol. 2 of Proc. Ent. Soc. Phila. 
was received at 9 May 1864 meeting. 

9lndex Scient. Cont. Jour. & Proc. Acad. Nat. Sci. Phila., 1812-1912: pp. xii-xiii. Gives dates of 
receipt for various parts of the early volume of the Proceedings. 

1 OTome I title page signature date qualified by preface date (p. xii). 

' 1 Trans. Ent. Soc. London 1869; p. xix (in Proc.). Notes receipt of Tomes 1-12 of the Ann. 
Soc. Ent. Belg. at 1 Nov. meeting. Tome 12 title page signature date: 1868-1869. I assume the 
date is fixed on the publication of Tome 12, I have no data for the separate except that its title 
page signature date is 1869. 



Journal of Research on the Lepidoptera 1(1) :73-83, 1962 

1140 W. Orange Grove Ave., Arcadia, California, U,S.A. 

Copyright 1962 


THE DISTRIBUTION OF THE SPECIES 
OF THE GENUS PIERIS IN NORTH AMERICA’ 

WILLIAM HOVANITZ = ’ 

1140 ]F. Orange Grove Ave., Arcadia, Calif. 


The North American species of the Genus Pieris are distributed 
from the Arctic Ocean to Guatemala. However, none of the species 
of the genus (sensu stricto) exist in habitats which would be con- 
sidered as tundra or arctic-alpine on the one hand, or as wholly tropical 
on the other. To this extent, the genus compares somewhat with Colias; 
the latter, however, contains two species that do exist in the tundra 
habitat. 

The concept of a species which has been applied here is the same 
as that which has been applied previously to the species of the genus 
Colias in drawing the distribution maps for that genus (Hovanitz, 
1950). This concept is based upon the consideration that the most 
important factor in stability of a species is the genetical population — 
or the interbreeding unit. Since, however, genes and their combinations 
cannot be seen, nor analyzed except by means of their effects on the 
appearance and physiology of the individuals carrying them, they must 
be studied through a comparative study of their morphological, anatom- 
ical or physiological characteristics. Thus, a name applied to one 
population may be applied to another population, or to many popula- 
tions, if their characteristics indicate that a similarity exists between 
the hereditary makeup of such different populations. Such a name 
may cover quite a diversity of differences in the same population, or 
even a diversity of different populations if they are genetically separated 
one from the other by geographical or ecological means. The extent of 
gene interchange as well as the evaluation of the isolation mechanism 
determines whether or not one population should be considered as the 

’Aided by a grant from tbe National Science Foundation, Washington, D.C. 

2The author wishes to thank the following for having made this distributional work possible 
by making available the collections under their care: C. B. MacNeil, California Academy of 
Sciences, San Francisco; L. M. Martin, Los Angeles County Museum, Los Angeles; T. N. Freeman, 
Entomological Research Institute, Ottawa; F. Rindge, American Mus'^um of Natural History, 
New York; Harry Clench, Carnegie Museum, Pittsburgh; W. S. Field and J. F. Gates Clark, 
U.S. National Museum, Washington, D.C.; and the Dept, of Entomology, Chicago Natural History 
Museum, Chicago, Illinois. 

3The base maps used in this paper are from Goodes Series of Base Maps by the University of 
Chicago Press, used by permission. 


73 


74 


HOVANITZ 


/. Res. tepid. 


’same species” as another, or a "subspecies” or "geographical race” 
of. it, or should not be considered significantly different. These criteria 
will be more exhaustively dealt with in a later paper. For practicability, 
different criteria must be used at different times and in varying circum- 
stances. The ultimate taxonomic designation is a product of weighing 
a great number of problems, some factors being known and others 
unknown, with the hope that the best result may be achieved. Another 
general rule underlies these studies; namely, that stability and uni- 
formity are desirable in nomenclature, and that past taxonomic 
decisions should not be upset by hasty, poorly-thought-out changes. But 
the past should not remain immutable when the decision to change is 
demanded by sound study or experimental evidence. Implicit in all 
taxonomic studies is perhaps an amount of individual opinion, perhaps 
greater than in many more experimental sciences, but even this can be 
greatly reduced by conscious effort to separate fact from opinion. It is 
the belief of the author that all facts upon which decisions are based 
should be stated; these are the "data” which can be refuted. New 
decisions should be made then only upon presentations of new data. 
Taxonomy by "hunches” is to be avoided; decisions should be 
accompanied by evidences for and against any decisions made. 

THE SPECIES OF PIERIS 

There are five clearly defined species of Pieris in North America. 
Of these, only four were native prior to 1800. Besides these, there 
are two other groups of related populations which might be considered 
"species,” or sibling species, and a large number of geographical varia- 
tions within each of these species, many of which are distinctive enough 
to be called "geographical races” or subspecies. Division of the species 
into geographical races is not to be considered in this paper, with the 
exception of the two sibling species just mentioned. 

Pieris rapae. This is one of two common European species of 
Pieris, the first immigrants* of which appear to have been introduced 
into North America prior to I860. Since that time, the species has 
spread over the whole of the Unitd States and much of Canada and 
Mexico, wherever the natural vegetation has been at least partially 
replaced by European weeds, such as black mustard, or by cruciferous 
agriculture. This species has a wide range of habitat which is suitable 
for its existence, since it is found from the warm parts of the Gulf of 
Mexico nearly to the tundra of Canada, and from the Imperial Valley 
below sea level to high mountain meadows in the Sierra Nevada or the 
Rocky Mountains. The food plants utilized are a variety of cruciferous 
plants, among which are all varieties of cruciferous garden vegetables 
(cabbage, kohlrabi, kale, etc.) as well as many weedy plants such as 
mustards, radish, sisymbrium, etc. Some garden flowers such as 
nasturtium are also utilized. In view of the "weedy” nature of this 
species, no map is given to illustrate its distribution. 


i(i): 73 -S 3 , 1962 


PIERIS DISTRIBUTION 


75 


Pieris napi (Fig. 1). This species is found throughout northern 
Europe and Asia where there is in effect a continuity with the forms 
of North America across the Bering straits and the islands of the 
Bering Sea, This is the most northern of all of the species of Pieris, 
its range extending to the edge of the tundra or slightly beyond along 
its northern limits. The northern distributional limit from the Macken- 
zie River and east follows closely the northern limits of the tree line. 
The species exists along both the Pacific and Atlantic coast lines, but 
extends farther southward on the Pacific Coast than on the Atlantic 
Coast. The southern-most population on the immediate Pacific Coast 
line is at Lopez Canyon, San Luis Obispo County, California at about 
35° north latitude. On the east coast at sea level, the most southern 
locality known is along the coast of New York and Connecticut 
opposite Long Island at 42° north latitude. However, at increasing 
elevations in the Appalachian Mountains southward, P. napi ( or 
P. vifginiensis) may be found as far south as the Great Smoky 
Mountains of Tennessee and North Carolina at about 35° north 
latitude. The southernmost extension of range of Pieris napi in the 
strict sense is in the vicinity of Connecticut and the Appalachians of 
Vermont, New Hampshire and Massachusetts. Westward, the range 
passes through the lower part of Ontario, Michigan, Wiconsin, Minne- 
sota, Manitoba, Saskatchewan and Alberta, always staying north of the 
prairies. A fuller discussion of the relation of Pieris napi to P. 
virginiensis is to be had in a separate paper to follow this one. 

Where the range of P. napi reaches the Rocky Mountains in central 
Alberta, the species extends clear to the Pacific coast and southward 
in the Rocky Mountains at increasing elevations to southern New 
Mexico and Arizona. The species appears to be absent in the drier 
ranges of the Great Basin and northward in the sagebrush country 
into south-central British Columbia. Elsewhere, it is widely distributed 
in the habitats that it prefers, these being damp wooded areas with 
partial shade, and temperatures not over seventy degrees Fahrenheit.. 

Of interest in the search for extensions of range are the following 
specific locations on the southern parts of the range of Pieris napi: 

California: Coast — - San Luis Obispo County, Lopez Canyon, 
December through May. 

California: Western side Sierra Nevada, Merced County, Yosemite 
Valley, 3-4000 feet, April-May. 

California: Eastern side Sierra Nevada, Placer County, 6-7000 feet, 
June. 

Nevada: No records at present. 

Arizona: Apache County, White Mountains, August; Gila County, 
"Globe” July. 

New Mexico: Otero County, James Canyon, Cloudcroft, August, 
Catron 6, Mogollon, May, August. 

Georgia: No records. 


76 


HOVANITZ 


/. Res, tepid. 



FIGURE 1. 


Map showing the North 
ndpi. 


American distribution of Pieris (Pieris) 


( i ):73-^3, 1962 


PIERIS DISTRIBUTION 


77 


South Carolina: No records. 

North Carolina: Black Mountains, April. 

Tennessee: Great Smoky National Park, Cherokee, April. 

Some other points of interest in the distribution of this species are 

the following records : 

Pribilof Islands, Alaska, July. Mouth of the Mackenzie River, North 
West Territories. 

Great Bear , Lake, N.W.T. Churchill, Manitoba ( Hudson Bay ) . 
James Bay, Hudson Bay, Quebec. Ramak, Labrador 

Pieris napi is the only known butterfly to inhabit the Aleutian 
Island^ or the islands of the Bering Sea. This ability is probably 
correlated with its adaptation to existence at temperatures below 70 °F, 
with low solar radiation and high humidity. 

Pieris (Pontia) protodice (Fig. 2). This is the most widely 
distributed species of Pieris in North America. Some prefer to place 
this species, together with the following two species, in a separate 
genus Pontia for which there is much to recommend. To do so, 
however, would reduce the size of the. genus Pieris beyond what would 
seem reasonable, and, therefore, to obviate this difficulty, and yet show 
phylogenetic relationship, Pontia would best be retained as a subgenus 
name. 

Pieris protodice is best adapted to habitats with more sunshine 
than is required by P. napi. It also is enabled to survive in habitats 
much too warm for that species. However, cold does not seem to be 
a limiting factor as long as there is a short warm season and plenty of 
sunlight. These conditions, to some degree, pertain to the area occupied 
by the species. It apparently does not flourish in areas of continual hot 
temperatures as indicated by its absence in all fully tropical climates. 

Pieris protodice differs from P. napi in its absence all along the 
immediate Pacific Coast from northern California through Alaska in 
keeping with its dislike for cool, cloudy areas. It is absent in the 
northern half of Alaska but eastward its distributional limits parallel 
those of the timbered area, as with P. napi. East of the Hudson Bay, 
however, P. protodice is absent north and east of Ottawa and New 
York State. The reasons for this event are baffling unless the present 
distribution of the species has been greatly disturbed by man in the 
eastern regions of North America as has been found true for Colias 
eurytheme. Pieris protodice prefers sunny, open fields as compared 
with P. napi which prefers partially shaded, damp areas. The cutting 
of the eastern deciduous forest to produce open fields for agriculture 
has aided both Colias eurytheme and Pieris protodice in extending their 
ranges and population density. It has probably a^so reduced the range 
and population density of Pieris napi since the habitats are conflicting. 
This probably accounts for the known diminution of Pieris napi in 
the region of Cambridge, Massachusetts where they were once exceed- 
ingy abundant, and now rare. 


78 


HOVANITZ 


/. Res. Lepid, 



FIGURE 2. Map showing the North American distribution of Pieris (Pontia) 
protodice. 


( 0 ! 73 -^ 3 , 


PIERIS DISTRIBUTION 


79 


Pieris protodice is found throughout the southeast from Long Island 
to the southern part of Florida and around the Gulf of Mexico to 
Guatemala. It is very abundant in the highlands of Mexico but may 
be found nearly everywhere except in woods or forested areas. It is 
found from the southern tip of Lower California northwards, having 
a great preference for open areas. It is possible that its great abundance 
in southern California coastal areas is a recent event, correlated with 
the change of the natural perennial grasslands to annual European 
grasses, mixed with Brassica nigra, the black mustard. It reaches its 
greatest abundance, however, in the desert regions where conditions 
of cool nights, hot days, full sunlight and periodic extensive abundance 
of food plants {Cleome, Brassica, Sisymbrium, etc.) allow the species 
to swarm in clouds. In such areas, the population abundance is usually 
limited by the periodicity of available water. 

In the mountain areas, the species is spread from the lowest valleys 
to the top of the highest mountains. The author has collected Pieris 
protodice at the top of Mt. Whitney, 14,500 feet elevation, the highest 
point in the United States south of Alaska, and in the bottom of 
Death Valley and Imperial Valley below sea level. 

The relationship of Pieris occidentalis to Pieris protodice is here 
left in doubt, but for the purposes of the map (Fig. 2), they are 
considered forms of one species. This point will be considered in more 
detail in a future paper. There is no doubt that at higher elevations 
and cooler temperatures, populations exist which have a blacker, fuller 
pattern on their wings than populations which exist at lower elevations 
at higher temperatures. The preponderance of the evidence suggests 
that these differences are not hereditarily controled, but that they are 
dependent entirely upon the environmental conditions. Within the 
species as a whole, as with P. napi, the darker indiyiduals exist in the 
more northern latitudes at cooler temperatures than the lighter ones. 
However, this relationship must be reconciled with local factors. In 
Colorado, Brown (1957) has shown that P. occidentalis appears to 
form definite populations at high elevations whereas P. protodice forms 
populations at low elevations. F. Rindge, however, in a personal com- 
munications has found both in considerable number in the same valley 
in Utah. These points cannot be reconciled at the present time. 
P. protodice when exposed to low temperatures develops a color 
pattern (more extensive melanin) which is similar to, or identical 
with, P, occidentalis. They could very likely be genetically the same 
and if they are genetically the same, they are not deserving of a scientific 
or Latin name of significance nomenclatorially. This point has not 
been proved, however. In the case of geographical variations, such 
deviations are considered subspecific unless proved otherwise. In the 
case of altitudinal or seasonal differences, such as in Pieris protodice 
or Colias eury theme, it is suggestd that Latin names not be applied 
except to geographically isolated populations and then in a subspecies 
sense only. 


80 


HOVANITZ 


/. Res. Lepid. 



FIGURE 3. Map showing the North American distribution of Pieris (Pontia) 
sisjmbrii. 


(i):73-h> 1962 


PIERIS DISTRIBUTION 


81 


Pieris (Pontia) sisymbrii. (Fig. 3) The Pontia subgenus of the 
Pierids all prefer dry habitats with considerable sunshine exposure, 
unlike Pieris napi or even Pieris rapae which prefer or can survive in 
shade and high humidity. Pieris sisymbrii is one of these. Its habitats 
are always open, highly exposed, usually rocky places. It prefers cool 
temperatures but with high solar radiation. It lives in the ’spring time 
in, the desert, as in the Mohave Desert where it flies at temperatures of 
highs not over 70° F, even though the temperatures later in the year 
will be over 100° F. It also lives in the mountains at elevations of over 
10,000 feet in June and July. In the coastal area of California, its 
habitats are usually dry, rocky areas (serpentine, etc.) which are more 
dry and more exposed to solar radiation than its general position on 
the map would indicate. 

Geographically (Fig. 3), the species exists throughout the moun- 
tainous areas of the west from the Yukon Territory to Mexico. It 
probably also exists in northern Mexico but there are no records. 
Outside of this area, there are a series of locations in the Northwest 
Territories from the Great Bear Lake southward past the Great Slave 
Lake to Lake Athabasca. Between these locations and the Rocky 
Mountains there are no locations known. This is probably due to lack 
of many suitable habitats. Collectors would do well to increase the 
known distribution of this species. It has a wide distribution but is 
characterized by the extreme isolated nature of them. Its habitats are 
narrowly restricted even on the desert and the adults have a short adult 
life, flying for only a few weeks during the year at any one place. The 
species aestivates and hibernates as a pupa which is very hard and 
impervious to water. 

There is relatively little variation between the individuals in a 
population in this species, and also relatively little variation geographi- 
cally between populations. Some variations have been described, how- 
ever, and will be considered in a subsequent paper. 

Pieris protodice is a species which is generally distributed and has 
many generations per year; it is therefore genetically and physiologi- 
cally pliable as is necessary to meet these changed conditions. On the 
other hand, Pieris sisymbrii is greatly restricted to isolated habitats, and 
the adults fly only a short time; thus, the species genetically may be 
said to have developed a narrower range of tolerance through a nar- 
rower range of selective factors of the environment. 

Pieris (Pontia) heckerii. (Fig. 4). Of all the species of North 
American Pieris, P, heckerii has the most restricted geographical range, 
extending from south-central British Columbia east of the Cascade 
Mountains southwards east of the Sierra Nevada into the deserts of 
southern California (Fig. 4). Its eastward limits coincide with the 
Rocky Mountain system in Montana, Wyoming and Colorado. The 
species does not extend into New Mexico so far as is known, unlike 
Pieris sisymbrii. The most southern-known limits are southwestern 


82 


HOVANITZ 


J. Res. tepid. 



FIGURE 4. Map showing the North American distribution of Pieris (Pontia) 
beckerii. 


(i):7}-Sl. 1962 


PIERIS DISTRIBUTION 


83 


Colorado, central Arizona, and southern Nevada, except for California 
where the species may be found in the foothills of the mountain ranges 
especially on the desert side. However, the species has been found on 
the coastal side in the area of the Santa Susana Valley (Moorpark), 
San Fernando Valley, Santa Clara River Valley and on the immediate 
coast from San Diego and southwards. 

Pieris heckerii exists at elevations of sea level at the coast and some 
parts of the desert to 9000 feet in the mountain valleys. It has always 
been found in areas that are relatively dry or desert-like with high solar 
radiation. Its distributional range coincides almost perfectly with that 
of the semi-arid brushlands extending from the rain-shadow valleys of 
central British Columbia, throughout the Great Basin and inter-moun- 
tain valleys of the Rocky Mountains. The factors that limit its distribu- 
tion in southern Arizona, New Mexico and northern Mexico are 
unknown. Perhaps there is a lack of proper food plant. The distribu- 
tion of the species in southern California south of the Great Basin 
is correlated closely with the distribution of Isomeris arhorea, its 
preferred food plant. It seems likely, however, that some other plant 
may also satisfy for this purpose. There is little doubt that intensive 
searching for additional poptilations of Pieris heckerii in the range 
of Isomeris will extend the distribution some considerable distance, 
especially in the south Coast Ranges of California, the foothills of the 
Sierra Nevada and southward in Baja California. 

Unlike Pieris sisymhrii which has a single short generation per 
year, in southern California, Pieris heckerii has a succession of genera- 
tions with the bulk of the adult flight being correlated with the time of 
of the winter rains. However, in the mountainous regions north of the 
Mohave Desert, the species has only one known generation, in the 
early summer. 

There is little geographical, local population, or seasonal variation 
in Pieris heckerii. However, some races have been described and will 
be considered in a later paper. 

LITERATURE CITED 

BROWN, F. M., D. EFF AND B. ROTGER. 1957. Colorado butterflies. 

Proc. Denver Mus. Nat. Hist. Nos. 3-7:1-368. 

HOVANITZ, WILLIAM. 1950. The Biology of Colias butterflies. 1. The 
distribution of the North American species. Wasmann J. Biol. 8:49-75. 


J 


Journal of Research on the Lepidoptera 1(1) :85-88, 1962 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

Copyright 1962 


DID THE CATERPILLAR EXTERMINATE 
THE GIANT REPTILE? 

S. E. FLANDERS 

University of California Citrus Research Center and 
Agricultural Experiment Station, Riverside, California 

The abrupt disappearance of the giant reptile during, the 
Cretaceous period brought to an end a very successful plant-animal 
system. For 120 million years the reptile population had roamed over 
large areas of the earth, subsisting in the lush vegetation covering such 
areas. From the standpoint of the reptile s subsistence, plant life was 
unlimited. It was an apparently inexhaustible resource. 

The possibility that the great reptiles disappeared as a result of 
malnutrition and starvation has been considered and discarded as quite 
untenable. That highly evolved plant-feeding animals do not normally 
become limited in numbers by starvation is axiomatic. Nicholson 
(1954) pointed out that the major influence which prevents phytopha- 
gous animals from reducing the earth s vegetation to extre'me sparseness 
is their attack by natural enemies. 

Exhaustibility of plant life, an inherent characteristic of many plant 
and animal relations, is rarely realized, being usually precluded through 
the regulation of the animal population by its natural enemies ( Flanders 
1959). Thus, the Cretaceous populations of the vegetarian reptiles, 
by serving as prey for populations of highly efficient predatory reptiles, 
were kept at levels well within the limits set by the amount of plant 
life. At the same time these predators served also to suppress the 
evolution of competitive types of animals of similar size such as the 
mammal. 

The geological record of the Cretaceous period, however, reveals a 
series of events which may have culminated in the temporary destruc- 
tion of vast am.ounts of plant life, a period in which the vegetarian 
reptiles may have been so devitalized that they either failed to 
reproduce or fell easy prey to their predatory relatives. 

The critical events which could have led up to the disappearance 
of the giant reptile were: (1) the replacement of the ancient fern- 
like plants by phanerogamic plants; and (2) the emergence of the 
Lepidoptera, an insect order which with few exceptions feeds only 
on such plants (Imms, 1930). 

The background against which these events should be visualized 


85 


86 


FLANDERS 


/. Res. Lepid. 


is the world-wide dominance of insects both in species and in popula- 
tion densities, a dominance which existed throughout the Age of 
Reptiles and the Age of Mammals. The pre-Cretaceous insect fauna 
appears to have been quite similar in kind and in numbers to the 
post-Cretaceous insect fauna except for the lack of species which fed 
on flowering plants (Carpenter 1952). A number of such species were 
subsequently derived from the pre-Cretaceous insect orders of Coleop- 
tera, Diptera, Hemiptera, Hymenoptera and Orthoptera, species whose 
immediate progenitors had presumably already acquired their natural 
enemies. 

According to Imms (1931), it is practically certain that the Lepi- 
doptera is pre-Tertiary in origin, highly organized individuals being 
represented in Eocene rocks. The paucity of insects in Cretaceous rocks 
is attributed to the absence in that period of suitable fresh-water 
deposits in which specimens could have accumulated and become 
fossilized. 

Plant-feeding insects like other plant-feeding animals acquire 
natural enemies which serve to conserve the plant life upon which they 
subsist. The fact that the conservation of plant life is a function of 
the natural enemies of Lepidoptera has been demonstrated during the 
past sixty years by the importation and establishment in North America 
of the natural enemies of the caterpillar for otherwise the reduction in 
brown-tail moth, Nygmia phaeorrhoea (Donovan), the satin moth, 
Stilpnotia salicis (Linnaeus), and the larch caseberer, Coleophora 
laricella (Hubner) (Clausen 1956). During the past three years 
the writer, by using the grain moth Anagasta and its natural parasite, 
Exidechthis, has demonstrated in the laboratory the role of a natural 
enemy in the conservation of plant material. In a constant environment 
parasitization continuously conserved the plant material despite the 
presence of a persistent feeding population of Anagasta. 

In nature, therefore, a community of plant-feeding animals con- 
sisting, say, of a pentatomid bug, a grasshopper, a moth caterpillar, 
a vole, a rabbit, and an ungulate depending on the same resource, grass- 
land vegetation, can remain stabilized for a long period of years ( Elton 
1946). Such stabilization, however, consists in the total amount of 
plant life conserved, not in the relative abundance of the plant species 
involved, an abundance determined by the phytophagous animals 
through their effect on the competitive capacities of the plant species 
(Wilson 1949). 

Devastations of plant life by caterpillars can occur today whenever 
the regulating action of their natural enemies is disrupted by environ- 
mental factors. Graham (1939) reported that during a ten-year period 
ending in 1920 an eruption of the spruce budworm, Choristoneura 
fumiferana (Clemens), was characterized by flights of moths so 
numerous that in the tree tops they had the apparance of a snow storm. 
During this eruption the caterpillars destroyed a volume of wood 


(i):85-88, 1962 


CATERPILLAR AND REPTILE 


87 


sufficient to supply all of the pulp mills then operating for a period 
of forty years. 

The capacity of a lepidopteran in the absence of its natural enemies 
to reduce great areas of lush vegetation to extreme sparseness and 
to maintain it so thereafter was demonstrated in Australia when the 
caterpillars of Cactoblastis cactorum (Berg) imported from Argentina 
in 1925 destroyed within six years approximately 50 million acres of 
the prickly pear, Opuntia spp. Dodd (1929) in reporting on this 
devastation stated that great care had been taken not to set free any 
of the natural enemies of the catrpillar for otherwise the reduction in 
prickly pear would have been of minor proportions. 

It is evident that the plant-consuming capacity of a caterpillar 
population could equal that of a giant reptile population and that this 
capacity because of the caterpillar’s very short life cycle could be 
attained at a much greater rate. Supplementing this capacity was a 
power of survival much greater than that of the giant reptile, the 
minimum food requirements of the individual caterpillar being 
infinitesimal relative to that of the individual reptile. 

Nevertheless, there is no reason to believe that the caterpillar and 
the giant reptile could not have subsisted together on the same 
vegetation if the caterpillar, immediately upon its Cretaceous 
emergence, had been subject to a regulation by natural enemies as 
effective as was that of the vegetarian reptile by its predatory relatives. 

The theory that the caterpillar exterminated the giant reptile rests 
on the assumption that between the emergence of the new insect order 
and its accessions of regulative natural enemies there occurred a brief 
period in which the caterpillar population regulated the world’s supply 
of plant food, a condition conducive to great variations in abundance 
of plant life and of caterpillar life, a condition characterized by "feast 
and famine" in which the caterpillar was able to survive but not the 
great reptile. 

The inherent weakness of the reptile was an extraordinary need 
for an abundance of plant material. Only a few years of plant scarcity 
could have exterminated it. Hordes of caterpillars, in rapid consump- 
tion of fig and breadfruit, laurel and willow, oak and magnolia, could 
have so restricted the spatial distribution of the giant reptile that 
either its diet was inadequate or it was unable to avoid its natural 
enemies. 

The small size of today’s descendent reptiles, the vegetarian turtle, 
the predatory crocodile, the snake, and the lizard, is evidence of the 
giant reptile’s elimination by starvation and predation. 

The abrupt end of the Age of Reptiles during the Cretaceous period 
is ascribed to a newly emerged order of insects, the Lepidoptera, on 
the supposition that for a brief period it regulated the world’s supply 
of plant life at starvation levels for the dependent reptiles. Thus, the 
giant reptiles which had survived during eons characterized by great 


88 


FLANDERS 


/. Res. tepid. 


changes in climate, continental uplifts, and different diets may have 

been exterminated by the lowly caterpillar. 

LITERATURE CITED 

CARPENTER, F. M. 1952. Fossil insects. 1952 U.S.D.A. Year Book. 
pp. 14-19. 

CLAUSEN, C. P. 1956. Biological control of insect pests in the continental 
United States. U.S.D.A. Tech. Bull. No. 113, 151 pp. 

DODD, ALAN P. 1929. The progress of biological centrol of prickly pear 
in Australia. Commonwealth Prickly-Pear Board, Brisbane Qld. 44 pp. 

ELTON, CHARLES. 1946. Competition and the structure of ecological 
communities. Jour. Animal Ecol. 15: 54-68. 

FLANDERS, S. E. 1959. Biological control. Jour. Econ. Ent. 52: 784-5. 

GRAHAM, S. A. 1939. Principles of forest entomology. McGraw-Hill, N.Y. 
410 pp. 

IMMS, A. D. 1930. A general textbook of entomology. E. P. Dutton & Co., 
Inc., N.Y. 703 pp. 

IMMS, A. D. 1931. Recent advances in entomology. P. Blakiston’s Son & 
Co., Inc., Philadelphia. 374 pp. 

NICHOLSON, A. J. 1954, An outline of the dynamics of animal popula- 
tions. Australian Jour. Zool. 2:9-65. 

WILSON, F. 1949. The entomological control of weeds. Int. U. Biol. Sci. 
Ser.B. 5:53-64. 


Journal of Research on the Lepidoptera 


1 ( 1 ) : 89 - 93, 1962 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
Copyright 1962 


FURTHER EVIDENCE OE THE DISTRIBUTION OF 
SOME BOREAL LEPIDOPTERA IN THE 
SIERRA NEVADA' 

C. H. ERIKSEN 

Los Angeles State College, Los Angeles, California 

Except where roads penetrate the high country, little or only 
scanty information is abaiiable concerning the animal life of the high 
Sierra of California. This condition is extant because these mountains 
are high, rugged and largely accessible only by trail. The fact also that 
collecting gear must be packed in and out encourages few people to 
undertake such studies. 

Our knowledge of the Lepidoptera fauna of the Sierra stems largely 
from the work of Garth (1935) and Tilden (1959). Garth’s study of 
the Butterflies of Yosemite National Park is notable since he not only 
provides a list of species taken within the confines of the park, but 
relates them to the life-zones in which they are normally found to fly. 
Tilden has refined the latter concept in his Tioga Pass studies by listing 
associations of smaller scope than life-zones. 

While backpacking the John Muir TraiP during the summers of 
1953, 1954 and 1955, the author made spot collections of Lepidoptera 
at various locations along the route (Figure 1). Such was a brief 
attempt to add information similar to that of Garth and Tilden to our 
knowledge of the day-flying Lepidoptera in high areas from Yosemite 
on the north to Mount Whitney on the south. All forms collected and 
altimdes at which they were taken are listed in Table 1. To alleviate 
any confusion, all nomenclature is after McDunnough (1938). 
Altitudes were determined from Starr ( 1953) . 

Since Garth (1935) and Tilden (1959) have shown that various 
species fly only within certain altitudinal ranges while others are 
unrestricted in their flight, there is little need to repeat similar fiindings 
here. However, the data collected does extend the known altitudinal 
range of several species. The following were taken at elevations higher 
than previously recorded and are associated with the life-zone of this 
extension. 

1 My thanks to Nelson Baker, Santa Barbara Museum of Natural History, for help with identifica- 
tion of the material and W. Hovanitz for aid in preparation of this paper. 

2The John Muir Trail follows a 215 mile route along the Sierra crest from Yosemite Valley on 
the north to Mount Whitney on the south. Except for a short distance out of Yosemite Valley 
(elevation 4,000'), altitudes range from approximately 7,000' to 14,I00'. 

89 


90 


ERIKSEN 


/. Res. tepid. 


Argynnis mormonia (Arctic- Alpine) 

Euphydryas chalcedona ( Transition, Canadian ) 

Lycaena helloides ( Canadian, Hudsonian ) 

Plebeius aquilo podarce ( Arctic-Alpine) 

Plebeius saepiolus ( Arctic-Alpine ) 

Plebeius icarioides (Hudsonian, Arctic-Alpine) 

Plebeius acmon ( Hudsonian ) 

Garth does not list Lycaena helloides above the Transition zone nor 
Plebeius saepiolus above the Hudsonian. Even though Tilden does not 
actually say, he intimates that both species transcend all zones. In the 
event that any confusion may arise, both species are included in the 
above list. 

It is of interest that those forms listed in this paper, which also 
are found in Colorado, have already been collected in that state from 
life-zones here described as extensions (Brown et al, 1957). 

LITERATURE CITED 

BROWN, F. M., D. Eff and B. Rotger. 1957. Colorado Butterflies. Proc. 

Denver Mus. Nat. Hist., Nos. 3-7:1-368. 

GARTH, JOHN S. 1935. Butterflies of Yosemite National Park. Bull. So. 
Cal. Acad. Sci., 34(1) :l-39. 

McDUNNOUGH, J. 1938. Check List of the Lepidoptera of Canada and the 
United States of America: Part L Macrolepidoptera. Mem. So. Cal. Acad. 
Sci., 1:1-275. 

STARR, JR., WALTER A. 1953. Guide to the John Muir Trail and the 
High Sierra Region. Sierra Club, San Francisco, 130 pp. 

TILDEN, J. W. 1959. The Butterfly Associations of Tioga Pass. Wasmann 
J. Biol., 17(2) :249-271. 


I (i): 8^-93, 19^2 


BOREAL LEPIDOPTERA 


91 



92 


ERIKSEN 


/. Re$. Lepid. 


Arctic Alpine 

Mather Plateau 

11,500* ; 7-30-53 

ro 





€n 












Bighorn Plateau 

11,500* 1 7-31-54 

bH' 




ffH 



a- 










Tyndall Creek 

11,000* i 7-29-54 

pH 





cO 


- 










Upper Bubbs Cr, 

11,000’ ; 7-29-54 






sH 












Evolution Lake 

10,990* s 7-26-53 

a> 

















Hudsonian 

Mdl* Palisade Low, L, 
10,650* t 7-28-53 

sH 

















Cascade Valley 

10,400* 1 7-5-54 


















Duck Uke Trail 

10,100* s 7-5-54 




- 










tH 




Purple Lake 

10,000* S 7-6-54 

CD 













ffH 



OI 

1000-Island L. 

9,850* 1 6-30-54 

OI 

■H 





(tH 











Little Pete Mdw, 

9,100* s 7-12-55 

O 

*H 

















Shadow Creek 

9,000* s 7-1-54 

(O 








(H 







CM 


Shadow Creek 

8,750* 1 7-2-54 


















Tuolumne Mdws, 

8,700* j 6-29-54 





CM 






pH 







Canadian 

Palisade Creek 

8,125* 1 7-31-55 















eH 

CO 


Quail Meadows 

7,700* 1 7-8-54 

CO 


iH 

tH 











.H 

ra 



Table 1. 

John Muir Trail Collection 
Sites, Listed by Elevation, 
and the Species and Number 
Taken 

Map Location Number 1 

Papilio zelicaon Luc, 

Papilio rutulus Luc, 

Parnassius clodius Men, 

> 

-o 

m 

m 

Q 

X 

>. 

s, 

p 

« 

m 

O 

o 

Colias behrii Edw, 

Pieris sisynbrii Bdv, 

Pieris occidentalis Reak, 

Argynnis montivaga Behr 

> 

(0 

•H 

G 

0 

1 
o 
s 

m 

G 

G 

< 

Brenthis epithore Edw, 

c 

o 

•o 

0 

0 

»H 

ns 

^ • 
o » 

m X 
r@ 

$4 

^ * 

>, > 

3 ^ 
U Q 

Euphydryas sierra Wgt, 

« 

m 

c 

•H 

a 

c 

>* 

o. 

3 

M 

Melitaea palla Bdv, 

Phyciodes laontana Behr 

Polygonia zephyrus Edw, 


(i); 8 ^- 93 . 1962 


BOREAL LEPIDOPTERA 


93 






























CO 




CM 

•H 



CM 








to 




CO 




<0 

r-t 



pH 



CO 













CO 














- 
































■H 



















•H 












- 














CM 










tn 


eH 














- 






pH 










•H 



CO 

•H 










































- 









•H 









CM 


•H 




- 







rH 








u> 






- 












CM 

CM 

CO 










Nymphalis antiopa L. 

Junonia coenia Hbn. 

Basilarchia lorquini Bdv. 

Callophrys dumetorum 

Derulexa 3. & B. 

Lycaena edit ha Mead 

Lycaena helloides Bdv. 

lycaena cupreus Edw, 

Leptotes marina Reak. 

Plebeius aquilo podarce 

F. £ F. 

Plebeius saepiolus Bdv. 

Plebeius icarioides Bdv. 

Plebeius acmon West & Hew. 

Plebeius shasta comstocki 
_E22$ ^ ^ 

Philotes battoides Behr 

Lycaenopsis pseudargiolus 
echo Edw. 

Thoybes nevada Scud, 

Polites sabuleti tecumseh 
Grin. 

Celerio lineata Fabr. 

Pseudohazis hera Harr. 

Anarta cordigera Thun. 



1 ( 1 ) : 95 - 96, 1962 


Journal of Research on the Lepidoptera 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
Copyright 1962 


ARGYNNIS AND SPEYERIA 

BY WILLIAM HOVANITZ 

Department of Zoology ^ Los Angeles State College 

The use of names for nomenciatorial purposes involves judgment 
on two levels. The first level involves nomenciatorial laws, and the 
second is the judgement on the part of the taxonomist as to the most 
reasonable and useful application of a name for a biological category. 

When a name is being considered for use on the generic level, 
the first point is its nomenciatorial availability. Here, it is important 
that the name not be a homonym, and that it have priority according 
to the established laws of nomenclature. A species is designated as the 
type species of that genus. For example, the species paphia L. has been 
designated as the type species of the genus Argynnis. 

Once the type species has been established for a genus, and there 
are no doubts of the nomenciatorial status of either, the specific name 
or the generic name, no one who wishes to remain within established 
criteria for nomenciatorial uniformity and fairness has any right to 
change this name. The generic name is strictly applied, however, only 
to the one species. It is the prerogative of any individual to use the 
same generic name for other species if he deems that the species 
concerned ought to be considered congeneric. Strictly speaking, it is 
possible for each species in existence to be the type species for a 
different generic name. If each of these names were used, each species 
would be in a different genus and the beneficial effects of binomial 
nomenclature would be rendered worthless. 

The purpose of a binomial nomenclature is to indicate relationships 
by grouping related species into the same genus. How this should be 
done is the prerogative of the individual taxonomist. It is assumed 
that such a person will use the method that is best suited to his purposes 
in showing relationships. 

The genus Argynnis in the broad sense is composed of many 
species, some of which can be grouped into categories having biological 
similarities, but which are smaller in content than the genus 
when considered with a broad view. Generic names have been used 
for some of these smaller categories with various species designated as 
types. It is perfectly satisfactory from a nomenciatorial as well as a 
biological standpoint to do this. This is the situation with regard to 
the use of Speyeria. The type species of the name as a genus is idalia. 


95 


96 


HOVANITZ 


/. Res. tepid. 


For those persons who wish to consider idalia as not congeneric with 
paphia, or with any other older name, Speyeria is a valid generic name. 
Or, it can be used in a subgeneric sense, in which case Argynnis would 
again be used as the generic name. 

This author believes that the broad use of the genus Argynnis is 
preferable to the practice of restricting the name to the paphia group 
of species, and further believes that the genus can be divided into some 
more or less satisfactory subgenera of which Speyeria is one. Our 
American species then would be designated Argynnis {Speyeria) idalia 
to show its relationship. The other American Argynnids then would 
also continue to use the name Argynnis rather than Speyeria. 

It is not the purpose of the editor of this journal to insist on any 
particular terminology for to do so would impinge on the freedom 
of the scientific worker, who alone has the right to make his choice. 
This editor can only insist on sound data to back the decisions of 
authors in their use of terminology, as in drawing conclusions from 
their work. 


advertisement 


TAKE CHLOROCRESOL ON YOUR NEXT TRIP 


Since the publication of the Chlorocresol method, 
field collecting techniques have been changed radi- 
cally. The Chlorocresol method eliminates the need 
for time consuming field mounting and/or relaxing 
of specimens. With Chlorocresol, specimens are re- 
tained in a relaxed condition for an unlimited period, 
allowing mounting to be done at a more convenient 
time or place. Field mounted specimens which would 
normally occupy a standard insect box may be packed 
in a container only 5" x 5" x IVi'- With specimens 
in a relaxed state, damage is minimized in the event 


of rough handling. 

Catalog No. 182 

Chlorocresol, 50 grams $0.95 

Chlorocresol, 100 grams 1.75 

Chlorocresol, 200 grams 3 00 

Styrene box, 5" x 5" x 114" per dozen 3.00 


Complete instructions are included with each 
shipment. 

Prices are F.O.B. Santa Monica, California. California 
residents please add 4% sales tax. 



BIO METAL ASSOCIATES 

Box 61 

Santa Monica, California 


Volume 1 


Number 1 


August^ 1962 



IN THIS ISSUE 


Editorial 1 

Variation in the silvering of Argynnis (Speyeria) callippe in 
the interior mountain area of south central California . . . 

Oscar Elton Sette 3 

The effect of various food plants on survival and growth rate of 
Pieris .... William Hovanitz and Vincent C. S. Chang 21 

General characteristics of the movements of Vanessa cardui (L.) 

J. W. Tilden 43 

Three factors affecting larval choice of food plant 

William Hovanitz and Vincent C. S. Chang 51 

The generic, specific and lower category names of the Nearctic 
butterflies. Part 1 — The genus Pieris . . Paddy McHenry 63 

The distribution of the species of the genus Pieris in North 
America William Hovanitz 73 

Did the caterpillar exterminate the giant reptile? .... 

S. E. Flanders 85 

Further evidence of the distribution of some boreal Lepidoptera 
in the Sierra Nevada Clyde Eriksen 89 

Argynnis and Speyeria William Hovanitz 94 

FRONT COVER ILLUSTRATION 

PARALLEL COLOR VARIATION IN SOME CALIFORNIA NYMPHALIDS 
Column 1, E.chalcedona 
Column 2, E.editha 
Column 3, C.leanira 
Column 4, C. pal la 
Column 5, C.hoffmanm 

Details in Ecology 22: 259-284, 1941. 





Ci> 




E 

@F RilSEARCHJ 
THE LEFI 




Volume 1 


Number 2 


]a^ary, 1963 




Established in 1 962 
Published at 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

Edited by WILLIAM HOVANITZ 

WITH EMPHASIS ON ENVIRONMENTALLY AND GENE- 
TICALLY INDUCED VARIATION, population analysis, evolution, 
phylogenetic taxonomy, zoogeography, comparative morphology, ecol- 
ogy, geographical variation, speciation, physiology, etc. In short, 
qimlity ivork on any aspect of research on the Lepidoptera. 

THE PURPOSE OE THE JOURNAL is to combine in one source 
the work in this field for the aid of students of this group of insects 
in a way not at present available. The JOURNAL will attempt to 
publish primarily only critical and complete papers of an analytical 
nature, though there will be a limited section devoted to shorter 
papers and notes. 

RATES: $ 8.00 per volume, personal subscription. 

$12.00 per volume, institutional subscription. 

All amounts are in U.S. dollars, payable in the U.S.A. 

AUTHORS ARE REQUESTED to refer to the Journal as an 
example of the form to be used in preparing their manuscripts. Fifty 
separates will be supplied to authors free; reprints will be sold at 
printer’s rates if ordered at time galley proofs are returned. If proofs 
are not returned promptly, the editor reserves the right to withhold 
publication, or to proceed with no responsibility. All issues of the 
Journal will be copyrighted; in submitting a paper each author agrees 
that the material has not been published elsewhere. 

MANUSCRIPTS AND SUBSCRIPTIONS should be mailed to the 
address noted above. 

THE LEPIDOPTERA FOUNDATION 

To provide permanent security to the JOURNAL, THE LEPIDOP- 
TERA FOUNDATION is being set up. Memberships are now being 
accepted at TEN DOLLARS per year, eight dollars of which goes to 
your subscription and two dollars goes into a permanent capital fund, 
the earnings only of which may be used for current publishing expenses. 
In addition a scale of higher contributions is to be set up which will 
be announced later. All funds received in excess of the subscription rate 
will go into the permanent fund. LIFE MEMBERSHIPS are offered 
now at Two hundred fifty dollars; these include the Journal for life. 
INSTITUTIONAL AND COMMERCIAL DONATIONS AND MEM- 
BERSHIPS are also available; rates will be announced later, or on request. 
These contributions are deductible. HELP YOUR JOURNAL. Meetings 
of members may be held at times and places to be arranged. 


Journal of Research on the Lepidoptera 1(2) :97-108, 1963 

1140 ’IF. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 196 } 


COMPOSITION AND RELATIVE ABUNDANCE 
IN A TEMPERATE ZONE BUTTERFLY FALMA 

THOMAS C. EMMEL and JOHN F. EMMEL 

Reed College, Portland, Oregon md Stanford University, California 

The Conner Pass area, lying on the crest of the Sierra Nevada 
range in Placer County, California, provides a rich variety of habitats 
lying between 6900 and 8300 feet elevation. During the summers of 
1956 and i 960 , the authors made an intensive study of this region and 
we have reported elsewhere (Emmel & Emmel, 1962a, 1962b) on 
the ecology and factors affecting distribution of the 74 species com- 
posing the butterfly fauna. We wish to report here the interesting data 
obtained on the faunal composition and relative abundance of species 
within the seven major Rhopalocera groups recorded in the Donner 
Pass area, and to evaluate the possible factors influencing this distribu- 
tion of species. 

INTRODUCTION 

Hovanitz (1958) has. recorded the occurrence of Rhopalocera 
families and genera at each five degrees of latitude and 1000 meters 
of elevation above sea level for the entire New World. However, this 
study did not include the numbers of species or genera in each family 
that occurred in each of these areas. Tilden (1959) divided the butter- 
flies of the Tioga Pass area under five plant associations; Emmel & 
Emmel (table 1, 1962a) partitioned the Donner Pass fauna under 
four plant associations. But these authors did not compare the faunas 
of each association or attempt to ascribe reasons for the greater number 
of species of a particular family in one association than another. The 
purpose of this paper is to report an examination of the composition 
and habitat associations of the Donner Pass fauna and from this 
examination to derive some understanding of the factors affecting the 
distribution and success of species in a butterfly family within a 
given set of environments. 

DESCRIPTION OF STUDY AREA & METHODS 

In a previous paper (Emmel & Emmel, 1962a) ^ four general 
vegetational associations representing the union of floras from three 


97 


98 


EMMEL AND EMMEL 


/. Res. Lepid. 


life zones (Transition, Canadian, and Hudsonian) were delineated, 
and may be briefly reviewed. 

1. The W et-M.eadow habitats (elevation 7000-7260 feet) support 
verdant expanses of grasses (Poa species) and sedges (Car ex), Lilium, 
Mimulus., Delphinium, Castilleia, and other wildflowers, with scattered 
thickets of willows (Salix) along the stream banks. Trifolium, Viola, 
Aster, Potentilla, and other lepidopterous food plants (see table IV) 
are common. 

2. Dry-Meadoiu areas (elevation 6800-7600 feet) are moist in 
the spring but are dry throughout July and August. Fireweed (Epilo- 
bium angustifolium) , Achillia, Calyptridium, and Penstemon, as well 
as grasses, are typical plants. Food plants include Kibes, Polygonum, 
Lupinus, Gnaphalium, and others as listed in table IV. 

3. Dense Forest (elevation 6800-7600 feet), mainly of Canadian- 
Zone trees, covers about one-half of the total Pass area. Red Fir 
(Abies magnifca), Pinus contorta, P. monticola, and Mountain Hem- 
lock (Tsuga mertensiana) represent the majority of the forest trees. 
Only three known butterfly food-plant genera are found here: Pinus, 
Arceuthobium, and Ceanothus, 

4. The Montane areas (elevation 7160-8383 feet) are generally 
open talus slopes, where the dry, almost soilless environment discourages 
the growth of most trees. Included in the flora of these mountain peaks 
are Rabbit Brush (Chrysothamnus) , Sage Brush (Artemisia), Mule- 
Ears (Wyethia mollis), Dipinus, and Sedum species. Eriogonum, 
Astragalus, Potentilla, Ceanothus, and other food plants in Table IV, 
are widely distributed here. 

Thus the diversity of vegetational associations in this temperate- 
zone locality provides habitats and food plants for a considerable 
number of butterfly species, and it is of interest to compare the relative 
success of species in different families in colonizing this favorable yet 
rather small area (six square miles) in the Sierra. In 1956 and I960, 
the number of species (see Emmel & Emmel, 1962a, for a detailed 
listing) in each of the four vegetational zones was determined by 
intensive collecting throughout Donner Pass. 

In the course of studying the influence of meteorological conditions 
on the flight activity of these species in I960, daily counts of the 
numbers of flying butterflies were made within the 500 x 800 feet 
"Lodge Meadow” study area (see Emmel & Emmel, 1962b), which 
contained 46 species recorded for the Pass (see Table III). This area 
is composed of both wet and dry meadows. These counts were made 
by direct observation and represented numbers seen for each species 
between 10 and 12 a.m. daily. The total number of butterflies re- 
corded in this latter study was 7,720. To determine "success” in 
numbers of flying adults, the daily counts for numbers of individuals 
for each of the species in a family were added together, and this figure 
was calculated as a percentage of the total for all families. Thus the 
contribution of each family toward the composition of the Donner 


(2):97-!oS, 196} 


DONNER PASS BUTTERFLIES 


99 


Pass butterfly fauna can be compared on a percentage basis, both as 
regards numbers of species and numbers of individuals. 

OBSERVATIONS 

Some 74 butterfly species were found in the Donner Pass area, 
and these species belonged to seven of the major families of Rhopalocera 
found in the western United States. The number of species in each of 
these groups varied from 1. fof the Danaidae to 28 for the Lycaenidae, 
or in terms of percentage of the total Pass fauna, 1.4% to 37.8% of 
ail the species. Table I shows the composition of the Rhopalocera groups 
recorded in the total Pass area. 

Table II and Figure 1 show the composition of the butterfly fauna 
recorded in each of the four general habitats of the Donner Pass area; 
the figures in Table II represent the per cent of total species in each 
habitat that belong to each family, while the graphs in Figure 1 show 
actual number of species for each family in each habitat. It is imme- 
diately obvious that the montane and dry-meadow habitats have the 
largest faunas of the four areas, and that the Lycaenidae family is usually 
dominant as regards number of species. 

As stated earlier, it was possible to obtain some data on the number 
of individuals of each species (46 in all) occurring in the Lodge 
Meadow study area. The composition and relative abundance of the 
six major Rhopalocera groups recorded there are given in Table III. 
For the most part, there is a good correlation between number of species 
and number of individuals for each family. 

DISCUSSION AND CONCLUSIONS 

The number of butterfly species found in this six-square-mile area 
of the Sierra Nevada is unusually high. Tilden (1959) records only 43 
species as occurring in the Tioga Pass region, representing a study area 
of approximately the same size although of somewhat higher elevations 
(over 9941 feet above sea level). Garth (1935) records about 100 
species for all of Yosemite National Park (area 1,179 square miles, 
elevations from 2000 to 13,090 feet), and the present authors and 
Lloyd M. Martin (personal communication) can note from their 
experience that the average restricted Sierran locality seems to support 
around 35 to 65 species. Thus the Donner Pass area is particularly 
interesting from the standpoint of invesigating the reasons for the 
occurrence and abundance of a species in a specific habitat, for as noted 
earlier (Emmei & Emmel, 1962a) this region is a meeting place for 
a variety of habitats- and a high number of plant species, and these 
factors may be viewed as possible reasons for this variety of Rhopa- 
locera. 

The relative proportions in number of species among the Rhopa- 
locera groups involved seems to follow the relative number as found 
throughout North America north of Mexico (determined from Ehrlich, 
1961 ). The descending order of these groups in terms of number of 


100 


EMMEL AND EMMEL 


/. Jl«, Lepid. 


TABLE I. Composition of the seven major Rhopalocera groups 
recorded in the Donner Pass area. 


Family 

No, of Species 

% of Total Species 

Papilionidae 

5 

6,8 

Pieridae 

8 

10,8 

Danaidae 

1 

1,4 

Satyridae 

2 

2.7 

Nymphalidae 

20 

27,0 

Lycaenidae 

28 

37,8 

Hesperiidae 

10 

13,5 

TOTALS 

74 - 

100,0% 


"In our earlier paper (Emmel 5 Emmel, 1962a) , 

74 species and 2 "forms”, Anthocaris sara reakirti and 
Colias eurytheme amphidusa, were listed, making a total 


of 76 phenotypically-distinct entities 




DONNER PASS BUTTERFLIES 


101 


species is (both in the continent and local Pass faunas): Lycaenidae, 
Nymphalidae, Hesperiidae, Pieridae, Papilionidae (follows Satyridae in 
the continent fauna), Satyridae, Danaidae. 

However, when the total Pass fauna is divided in terms of occurrence 
in each of four habitats, patterns of apparent preference for certain 
habitats emerge — both for all families combined and for individual 
groups (Table II and Figure 1). The highest total number of species 
(60) occur in the montane habitats, the next highest (55) in the dry 
meadow areas, the third highest (44) in the wet meadows, and the 
lowest number (26) in the forest areas. The descending sequence of 
environmental support of number of butterfly species is thus: 

Montane > Dry Meadows > Wet Meadows > Forest. 

This pattern, obtained by comparing numbers of Rhopalocera species, 
is probably explained by the observed fact that the montane areas 
contained a greater variety of ' mico-habitats” than any other area. These 
micro-habitats included small stream areas, exposed mountain tops, 
open and brushly talus slopes, etc., ranging from 7100 to 8383 feet in 
elevation, the greatest range in altitudes for any of the four "macro- 
habitat” areas considered. Such variation in ecological conditions 
(especially from edaphic and climatic standpoints) permits a wide 
variety of potential food plants to flourish. Axelrod (I960) has 
commented on the marked diversity of habitats in montane areas 
and the possible influence of this condition on early angiosperm evo- 
lution through permitting rapid development and adaptive radiation. 
Such diversity also influences butterfly distribution, variation, and 
speciation (see Emmel & Emmel, 1962a; Le Gare & Hovanitz, 1951; 
Moeck, 1957 ), and provides potential areas for successful invasion by 
butterflies already occurring elsewhere. 

The large number of total species in the dry meadow areas is 
believed to be due to the great variety of known food plants occurring 
there; the wet meadows supported correspondingly fewer food-plant 
species. The jorest areas contained the least variety of micro-habitats 
and food plants; also, solar radiation, an important influence on butter- 
fly flight habits (see Emmel & Emmel, 1962b), was obviously at a 
considerably lower level in this habitat than in the other three. Table 
IV shows the number of known food-plant genera in each habitat for 
each Rhopalocera group. These and the preceding data agree with the 
theory that a greater variety in each of these Sierran butterfly groups 
is more likely to occur where micro-habitats and food plants are most 
varied, as these conditions, together with a high level of solar radiation, 
provide support for a wider selection of species than in such areas as 
pine forests. 

This theory can be applied equally to the question of why one 
group has more species than another group in a certain habitat. The 
Papilionidae are more successful in forest than in montane areas in 
terms of per cent of total species there, but they actually have the 
most species in the latter areas. So we must consider: 


102 


EMMEL AND EMMEL 


/. Res. Lepid. 


WET MEADOWS DRY MEADOWS FOREST 


MOr^TANE 


4 



PAPiLION/DAE 


6 

4 

2 


PIERIBAE 


DANAIDAE 

Is;:; 


SATYRIDAE 





HESPERUDAE 


Fig. 1. Numerical composition of the butterfly fauna recorded io each of 
four general habitats of the Donner Pass area. 


the 


NUMBER OF SPECIES 



i (2):^7- io 8, 15163 


DONNER PASS BUTTERFLIES 


103 


1) That habitat in which the group has its highest percentage 
of the total species population: 

Papilionidae Forest {1.1%) 

Pieridae Forest (15.4%) 

Danaidae Forest (3.8%) 

Satyridae Montane (3.3%) 

Nymphalidae Forest (38.4%) 

Lycaenidae Wet Meadows (45.5%) 

Hesperiidae Forest (19.2%) 


2) That habitat in which 
number) : 

Papilionidae 

Pieridae 

Danaidae 

Satyridae 

Nymphalidae 

Lycaenidae 

Hesperiidae 


the group has the most species 
Montane (4) 

Dry Meadows, Montane ( 6 ) 
( 1 in each habitat) 

Montane (2) 

Montane (16) 

Dry Meadows ( 22 ) 
Montane (10) 


( direct 


The really significant consideration in evaluating a group’s contribu- 
tion to the fauna in a series of habitats would seem to be the number 
of species it has existing in each habitat. Of course, a further and 
perhaps more important consideration would be the number of indi- 
viduals of the group in that habitat, but these data were not obtained. 
However, speaking from the standpoint of number of species the 
butterfly groups are uniformly most successful in the montane and 
dry meadow habitats. 

As opposed to evaluating the most suitable habitat for most groups, 
one can also consider the success of individual groups in colonizing 
or existing in the various habitats. In the wet meadow, dry meadow, 
and montane habitats, the Lycaenidae are the most abundant group — • 
with 20 to 22 species in such areas. The Nymphalidae take second 
place (10 to 16 species) except in the forest zone where they lead the 
other groups in number of species. The Hesperiidae are third — with 
5 to 10 species in each zone — except for being second in the forest 
areas (leading the Lycaenidae). The Pieridae are uniformly fourth 
(4 to 6 species) while the Papilionidae have only 2 to 4 species in 
each zone. There is only one species in the Danaidae (all habitats, 
however) and two in the Satyridae (montane habitat only). 

This apparent relative success of each group, in our opinion, is not 
due to a "better” adaptation (the result of natural selection) of a 
group’s species in one or another habitat, as Ehrlich (1962) has 
emphasized. Instead, from our observations, success is likely due to 
the fact that food plants of species in the seven Rhopalocera families 
were more varied (as to number of kinds; Table IV), generally more 
numerous (as to number of individuals of each plant species), and 


104 


EMMEL AND EMMEL 


/. Res. Lepid. 


TABLE II. Composition of the butterfly fauna recorded in each of the 

j. 

four general habitats of the Donner Pass area. 


Family 


Habitat 

Area““ 



Wet Meadows 

Dry Meadows 

Forest 

Montane 

Papilionidae 

4.5 

5.5 

7.7 

6.7 

Pieridae 

9.1 

10.9 

15.4 

10,0 

Danaidae 

2.3 

1.8 

3.8 

1.7 

Satyridae 

0.0 

0.0 

0.0 

3,3 

Nymphalidae 

27.3 

27.3 

38.4 

26.7 

Lycaenidae 

45.5 

41.0 

15.4 

35.0 

Hesperiidae 

11,4 

14,5 

19.2 

16.7 

(1) TOTAL SPECIES 

44 

55 

26 

60 

(2) PERCENTAGE OF 
TOTAL PASS FAUNA 
(74 species) 
REPRESENTED: 

59.5% 

74.3% 

35.2% 

81.1% 


“The numbers in each vertical column (file) represent percent of 
total species in that one habitat (the total number of species is found 
at the bottom of each column). The actual numbers of species of that 
family in each habitat, from which the preceding percentages were 
calculated, are shown in graphic form in Figure 1, 

““See Emmel S Emmel (1962a) for map and complete description of 
all habitats in the Pass area. 


i(2);97-to8, 196} 


DONNER PASS BUTTERFLIES 


105 


more generally distributed for the Lycaenids than for the Nymphalids, 
for Nymphalids than for Hesperids, etc. It is concluded in this paper 
that the composition of a butterfly fauna is affected by the require- 
ments (or adaptation) of each butterfly species in that fauna for a 
certain food plant, in addition to a certain level of solar radiation and 
other factors as noted in Emmel & Emmel, 1962a. In turn, food plants 
obviously have grater chances of finding proper environmental condi- 
tions in a macro-habitat that contains a number of varied micro-habitats, 
each with particular edaphic and climatic conditions. The greater the 
number of host food plants, the greater the number of potentially 
successful butterfly species, and this theory is believed to account for 
the observed distribution of the 74 butterfly species (of seven major 
groups) among the four macro-habitats of the Donner Pass area. 

TABLE III, Composition and relative abundance of the six major Rhopalocera 
groups recorded in the Lodge Meadow habitat. 


Family 

No, of Species 

% of Total Species 

% of Total 
Individuals 
Recorded 

Papilionidae 

2 

4.4 

4.7 

Piaridae 

4 

8,7 

9,8 

Dana i da e 

1 

2.2 

0.3 

Nymphalidae 

15 

32,6 

44.9 

Lycaenidae 

16 

34.8 

23,2 

Hesperiidae 

8 

17.4 

16,7 

TOTALS 

46 

100.1% 

99 . 6 % 


SUMMARY 

1. The Donner Pass area in Placer County, California, supports 
an extraordinary number (74) of Sierran Rhopalocera species, and 
its varied habitats and vegetational associations make the area par- 
ticularly interesting for investigating the basis for the occurrence and 
abundance of a species and a family in a specific habitat. 

2. The number of species in the seven major families of the Pass 
area are proportionately equivalent to the numbers of species in these 


106 


EMMEL AND EMMEL 


/. Res, Lepid. 


TABLE IV, Number of known food plant genera in each habitat 


for each 


Rhopalocera family" 


Rhopalocera 


Habitat 

Area 


Family 

Wet Meadows 

Dry Meadows 

Forest 

Montane 


PAPILIONIDAE 

Salix 

— 

Ceanothus 

Cymopterus 

Salix (stream) 
Ceanothus 

Prunus 

Sedum 

FIERI DAE 

Cruciferae 

genera 

Cruciferae 

genera 

Pinus 

Cruciferae 

genera 


Trifolium 

Trifolium 



DANAIDAE 


- 

- 

- 

SATYRIDAE 

( Grasses ) 

(Grasses) 

(Grasses) 

Grasses 

NYMPH ALI DAE 

Viola 

Castilleia 

Salix 

r’lantago 

Castilleia 

Aster 

SlrsEum 

Ribes 

Gnaphalium 

Lupinus 

Ceanothus 

Castilleia 

Aster 

Chrysopsis 

Ceanothus 

Salix (stream) 
Lupinus 

LYCAENIDAE 

Salix 

tupinus 

Ceanothus 

Lupinus 

Arceuthobium 

Pinus 

Ceanothus 

Lupinus 


trifolium 

Potentilla 

Polygonum 

Ribes 

Eriogonum 

Potentilla 

Calyptridium 




Polygonum 

Sedum 

Eriogonum 

Astragalus 

HESPERIIDAE 

Potentilla 

Grasses 

Sidalcea? 

Potentilla 

Grasses 

Sidalcea 

— 

Potentilla 

Grasses 


"Data summarized from text of Eramel 6 Emmel, 1962a. 


(2):97-Jo8, 1963 


DONNER PASS BUTTERFLIES 


107 


groups as found for the whole of North America (north of Mexico). 
From most to least number of species: Lycaenidae, Nymphalidae, 
Hesperiidae, Pieridae, Papilionidae (follows Satyridae for North 
American fauna ) , Satyridae, Danaidae. 

3 . The highest total number of butterfly species in Donner Pass 
occurs in the montane habitats ( 60 species ) . The dry meadow habitats 
support 55 species, the ivet meadow habitats have 44 species, and the 
jorest areas have 26 species occurring in them. 

4. The basis of this distribution is explained by the theory that 
a greater food-plant diversity occurs in the areas (such as the montane 
macro-habitats) that have a number of micro-habitats, each with 
particular edaphic and micro-climatic conditions, and that this host- 
plant diversity promotes the immigration and continued successful 
existence of more Rhopalocera species than in less diversified macro- 
habitats ( such as these Sierran forest habitats ) . The possible influence 
of such factors as solar radiation and climate acting directly on butter- 
flies rather than plants is also considered. 

5 . Data obtained on the number of flying individuals of all species 
found in the Lodge Meadow area showed there was usually a good 
correlation between number of species and number of individuals for 
each family in this restricted Sierran locality. 


SYSTEMATIC LIST OF RHOPALOCERA OCCURRING 
AT DONNER PASS, PLACER COUNTY, CALIFORNIA 
PAPILIONIDAE 


1. Papilio zelicaon Lucas 

2. Popilio indra indr a Reakirt 

3 . Papilio rutulus Lucas 

4. Papilio eurymedon Lucas 

5. Parnassius clodius baldur 
Edwards 

PIERIDAE 

6. Neophasia menapia 
Felder & Felder 

7. Pieris sisymbrii Boisduval 

8 . Pieris protodice Linnaeus 

9 . Pieris rapae Linnaeus 

10. Euchloe creusa hy antis Edwards 

11. Anthocaris sara form julia 
Edwards 

12. Colias eurytheme Boisduval 

13. Colias phUodice eriphyle 

Edwards 

DANAIDAE 

14. Danaus plexippus Linnaeus 
SATYRIDAE 

15 . Ceononympha tullia calif ornica 
Westwood 

16 . Cercyonis sthenele oetus 
Boisduval 

NYMPHALIDAE 

17. Speyeria cyhele leto Behr 

18. Speyeria zerene zerene 
Boisduval 


19 . Speyeria coronis snyderi Skinner 

20. Speyeria atlantis irene Boisduval 

21. Speyeria mormonia arge Strecker 

22. Boloria epithore Edwards 

23 . Chlosyne palla Boisduval 

24. Chlosyne hoffmanni hoffmanni 
Behr 

25 . Phyciodes campestris montana 
Behr 

26 . Phyciodes mylitta Edwards 

27. Polygonia zephyrus Edwards 

28. Nymphalis calif ornica Boisduval 

29 . Nymphalis milberti Latreille 

30. Nymphalis antiop a Linnaeus 

31 . Vanessa cardui Linnaeus 

32 . Vanessa atalanta Linnaeus 

33 . Vanessa virginiensis Drury 

34 . Vanessa car ye Hubner 

35. Precis lavinia Cramer 

36 . Limenitis lorquini Boisduval 
& Leconte 

LYCAENIDAE 

37. Satyrium calif ornica Edwards 

38. Satyrium sylvinus Boisduval 

39. Satyrium saepium Boisduval 

40. Satyrium behrii Edwards 

41. Satyrium fuliginosa Edwards 

42. Strymon melinus Hubner 

43 . Callophrys johnsoni Skinner 

44. Callophrys nelsoni Boisduval 


108 


EMMEL AND EMMEL 


/. Res. Lepid. 


45. CaUophrys augustinus iroides 
Boisduval 

46. CaUophrys eryphon Boisduval 

47. CaUophrys dumetorum perplexa 
Barnes & Benjamin 

48. Lycaena arota virginiensis 
Edwards 

49. Lycaena edit ha Mead 

50. Lycaena nivalis Boisduval 

5 1 . Lycaena cupreus Edwards 

52. Lycaena heteronea Boisduval 

53. Everes comyntas amyntula 
Boisduval 

54. Plebejus anna Edwards 

55. Plebejus saepiolus Boisduval 

56. Plebejus icarioides Boisduval 

57. Plebejus shasta Edwards 

58. Plebejus acmon Westwood 
& Hewitson 

59. Plebejus {acmon?) lupini 
Boisduval 


60. Agriades glandon podarce 
Felder & Felder 

61. Glaucopsyche lygdamus hehrii 
Edwards 

62. Philotes enoptes Boisduval 

63. Philotes battoides intermedia 
Barnes & McDunnough 

64. Celastrina argiolus echo 
Edwards 

HESPERIIDAE 

65. Thorybes nevada Scudder 

66. Pyrgus ruralis Boisduval 

67. Pyrgus communis Grt. 

68. Erynnis juvenalis Fabricus 

69. Erynnis afranius Lint. 

70. Hesperia juba Scudder 

71. Hesperia nevada Scudder 

72. Hesperia harpalus Edwards 

73. Polites sonora Scudder 

74. Polites sabuleti tecumseh Grin. 


ACKNOWLEDGMENTS 


The authors again express their appreciation to Mr. William N. Goodall 
of the National Audubon Society, Director of the former Audubon Camp of 
California at Donner Pass (Norden, California), for extending full coopera- 
tion to us during the collection of the data reported in this series of three 
papers on the Donner Pass butterfly fauna. Drs. Thomas Harvey and Kenneth 
Tanksley of the Camp staff kindly furnished plant identifications. Discussions 
with Mr. Lloyd M. Martin, Associate Curator of Entomology at the Los 
Angeles County Museum, have illuminated a number of points included in 
these papers. The authors wish to thank Dr. G. Frank Gwilliam, Associate 
Professor of Biology at Reed College, for his help in preparing the present 
paper. 

LITERATURE CITED 


AXELROD, DANIEL 1. I960. The evolution of flowering plants. In The 

Evolution of Life, Sol Tax, ed. Pp. 227-305. Univ. of Chicago Press. 
EHRLICH, PAUL R., & A. H. 1961. How to Know the Butterflies. Brown, 
Dubuque, Iowa. 262 pp. 

— — — - — ■, & RICHARD W. HELM. 1962. Patterns and populations. 

Science 137: 652-657. 

EMMEL, THOMAS C., & JOHN F. EMMELL. 1962a. Ecological studies of 
Rhopalocera in a High Sierran community — Donner Pass, California. 

I. Butterfly associations and distributional factors. Journ. Lepid. Soc. 

16: 23-44. 

— — , 1962b. Ecological studies of Rhopalocera in a High 

Sierran community — Donner Pass, California. II. Meteorologic in- 
fluence on flight activity. Journ. Lepid. Soc., in press. 

GARTH, JOHN S. 1935. Butterflies of Yosemite National Park. Bull. 
So. Cal. Acad. Sci. 34: 37-75. 

HOVANITZ, WILLIAM. 1958. Distribution of butterflies in the New 
World. In Zoogeography. American Association for the Advancement 
of Science, Publication No. 51, Washington, D. C. Pp. 321-368. 

LE GARE, MARY JUDE, & WILLIAM HOVANITZ. 1951. Genetic and 
ecologic analyses of wild populations in Lepidoptera. 11. Color pattern 
variation in Melitaea chalcedona. Wasmann J. Biol. 9" 257-310. 

MOECK, ARTHUR H. 1957. Geographic variability in Speyeria. Mil- 
waukee Entomological Society. 48 pp. 

TILDEN, J. W. 1959. The butterfly associations of Tioga Pass. Wasmann 

J. Biol. 17: 249-271. 


Journal of Research on the Lepidoptera 1(2) : 109-1 13, 1963 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 1963 


THE ARGYNNIS POPULATIONS OF THE SAND 
CREEK AREA, KLAMATH CO., OREGON 

Part I: The effect of the formation of Mt, Mazama on the area 

AND ITS POSSSIBLE INFLUENCE ON THE BUTTERFLY FAUNAS OF THE 

Sand Creek Basin. 


J. W, TILDEN 

San Jose State College, San Jose, California 


INTRODUCTION 

The populations of Argynnis (Speyeria) found in eastern 
Oregon share with similar populations in many parts of western 
United States, a tendency to great variation and to a breakdown of 
the distinctive appearance of many of the named geographical pheno- 
types (subspecies). The region drained by Sand Creek, lying east of 
Crater Lake in Klamath County, Oregon, is occupied by populations 
of Argynnis exhibiting this variation to a marked degree. 

The Sand Creek area was chosen for study not only because of 
the variability of its Argynnis fauna, but in particular because it is 
a region the recent geological history which is fairly well understood, 
and inferences may be drawn of the possible effect of this geological 
history on the several species of Argynnis now inhabiting this region. 

Part I of this paper will deal with the recent geological history of 
Mt. Mazama and the nearby lands to the east of Crater Lake, and the 
possible influence of these events on the faunas now living in this 
region. Part II will attempt an analysis of the variation of these 
Argynnis populations, and is postponed until much more field work 
can be done so that a proper appreciation of the extent of variation 
is possible. 

BACKGROUND 

The stability of certain isolated populations has been interpreted 
as due to long occupation of the present habitat. This time element 
has been construed as allowing stabilization, without undue influence 
from surrounding populations. Such conditions infer stability in the 
habitat, without the disturbing influence of earth changes in the area. 
Such stable habitats have been referred to as refugia. 

Opposed to such a condition of stability, many portions of the 
earth s surface have been subjected to changes more frequently, and 
some of these changes have occurred in very recent times. Eastern 
Oregon is a region of many lava flows, and scoria and cinder falls, and 


109 


110 


TILDEN 


/. Res. Lepid. 


the effects of these actions are still clearly visible in numerous places. 
It is suggested that such phenomena must have had a major effect 
upon any butterfly populations existing in the regions where such 
events took place. 


THE SAND CREEK BASIN AND ITS 
RECENT GEOLOGICAL HISTORY 

Sand Creek rises at Anderson Spring, elevation 7088 ft, slightly 
northeast of Kerr Notch in Crater Lake National Park, and runs 
easterly, leaving the park in the near vicinity of the now unused East 
Entrance. It continues out of the park, the elevation gradually decreas- 
ing. The area chosen for study is that from Lost Creek Campground, 
Crater Lake National Park, 5972 ft., located between Sand Creek and 
its branch, Wheeler Creek, and the point at which Highway 232 
("Old 97”) crosses Sand Creek. This area is readily accessible. Work 
in the Crater Lake National Park was possible under a permit from the 
National Park Service. 

The Argynnis samples indicate that the following species are repre- 
sented: Argynnis {Speyeria) coronis Behr, zerene Bdv., callippe Bdv., 
egleis Behr, atlantis Edw., and hydaspe Bdv. Possibly others may be 
present in lesser numbers but have not yet been collected. It is inter- 
esting to note that Argynnis cybele {leto) appears to be absent from 
the region, and also from all of Crater Lake National Park, though 
common west of the Cascade divide, and also in the Blue Mountains 
of eastern Oregon. The scarcity in the Sand Creek Basin of suitable wet 
habitats, or the recency of earth surface changes, may be clues to its 
absence. 

Fair samples of these various species, collected by the author and 
also by others, have been examined, and it is evident that the degree 
of variation is surprising. As stated above, analysis of these and future 
samples is postponed until later, but enough information is now avail- 
able to make it apparent that the populations of each species involved 
are far from stable. Many individuals resemble no named geographic 
segregate. Some species are represented by individuals resembling two 
or more named segregates and it is evident that no equilibrium in 
phenotype has been reached by any of the several species inhabiting 
the area. 

Crater Lake is not a true crater, but a caldera, brought into its 
present form by the collapse inwardly of the summit of Mt. Mazama. 
In its most developed state, Mt. Mazama was more than 12,000 ft. in 
height. During the Pleistocene, it developed its greatest elevation, 
produced a great lava flow, and was subsequently glaciated extensively. 
The effects of this glaciation are visible around the present rim of the 
lake. Later, the present Crater Lake was formed by the collapse inwardly 


i(2):io^-Jij, 196} 


SAND CREEK ARGYNNIS 


111 


of the upper mile or so of Mt. Mazama, with attendant glowing 
avalanches of scoria and pumice and great clouds of pumice dust. 
The area east of the lake was covered with pumice and scoria from 
the glowing avalanches. The land still further east was covered by 
pumice falls that reached a depth of ten feet or more close to the 
mountain, and gradually thinned out easterly, extending in at least a 
very thin layer, approximately fifty miles to the east. 

It is believed that the glaciers, or some of them, must still have 
been present when the collapse took place, since the residue of scoria 
and pumice is light in certain areas around the rim. This would be the 
situation if the scoria and pumice fell onto glaciers and was carried 
away in part. If it had fallen in every case on bare ground, it would 
be left there. This may explain why the present pumice soils are more 
extensive in the non-glacial areas. It is believed also that the collapse 
took place in winter, because there is relatively little evidence of 
severe forest fires in the wake of the collapse. If the collapse had taken 
place in the summer, it has been reasoned that the forest fires surely 
would have been more extensive. 

There seems to be agreement that most or all of the vegetation was 
destroyed within the effective range of the scoria and pumice avalanches 
and of the heavier pumice fall. If this is correct^ the present vegetation 
of the entire region must have regrown since the collapse took place. 
The rate of recovery of the vegetation seems to have been inversely 
proportional to the depth of the pumice and scoria that covered a 
given area. The prevailing winds are believed to have been much the 
samx then as now, basically western, since the pumice was carried much 
further east than west. 

The Sand Creek Basin is situated so that it received very heavy 
pumice fall, and very probably, extensive to complete vegetational 
destruction. 

Carbon dating of wood that is imbedded in the pumice, and which 
is therefore assumed to have been charred by the glowing avalanches 
of pumice and scoria, sets the time of the collapse at 6640 ± 250 years. 
In round numbers, popular accounts state this as 6550 years. From 
this it is inferred that the Sand Creek Basin became a sterile habitat 
at that time. 

It is of interest to observe that such streams as Upper Sand Creek 
and Wheeler Creek, below Kerr Notch, have deep V-shaped canyons, 
geologically very young and without sign of glaciation such as that 
found on the rocks of the Rim above. From this it seems clear that 
Sand Creek acquired its present bed recently. Indeed, its present course 
may very well be consequent to the recent geological events of the 
area. The depth of the pumice is also clearly visible. 

Knowledge of the date when the vegetation was destroyed allows 
some speculation concerning present vegetational cover of the region. 
Recovery of the vegetation must have lagged for many years. The 
types of vegetation found in the total Crater Lake region permit some 


112 


TILDEN 


/. Res. Lepid. 


understanding of the relative times of recovery. In Crater Lake National 
Park, the forest along the southern boundary is a mature Sierran Coni- 
ferous Forest of Ponderosa Pine, Sugar Pine, White Fir, Douglas Fir 
and Incense Cedar, with an understory of Snow Brush {Ceanothus 
velutinus) and Greenleaf Manzanita {Arctostaphylos patula) . The 
forest of the flanks of the mountain on the southern and western slopes 
is a mature Mountain Hemlock-Shasta Fir Forest, with increasing 
amounts of Subalpine Fir at higher elevations. Such forests have 
relatively less understory. The highest elevations support Whitebark 
Pine. 

Northerly and easterly, in the areas of greater pumice fail, (as for 
instance the Pumice desert) the forest has not even yet succeeded in 
taking over the denuded and pumice-covered land. In the intermediate 
areas where the forest is slowly invading and colonizing the .pumice 
soils, the successful pioneer is Lodgepole Pine, which on such soils 
forms a singularly uniform and barren type of forest with very few 
underplants. 

The soils of the study area are formed of pumice and scoria and 
from the foregoing evidence, are derived from the pumice fall from 
the collapse of Mt. Mazama. The forest here is essentially Lodgepole 
Pine Forest, of the type found also on the pumice at higher elevations 
in the park. The soils and the forest are evidently contemporary in the 
Sand Creek Basin, with those occurring on pumice within the park 
itself. We should regard the forest cover of the Sand Creek Basin as 
equally recent. 

Dating of the pumice fall and the destruction of the vegetation 
give a reference point. Vegetational recovery must involve a very 
considerable period of time. This reasonably will be longer in areas 
where the pumice fall is deeper. While no exact figure can at present 
be suggested for the age of the current forest cover of the Sand Creek 
Basin, it must be very much less than 6640 ± 250 years. Very likely 
portions of it may be a fraction of this age. 

From this it seems evident that the Argynnis populations of the 
Sand Creek Basin (and similar areas) are occupying habitats that 
are the opposite of refugia. The time in occupation of contemporary 
populations on such disturbed areas is brief. 

DISCUSSION 

Assuming an area from which vegetation has been destroyed, with 
attendant soil changes, and with sufficient time elapsed to allow new 
vegetation to occupy the area, we may well expect certain things to 
take place, among which the folowing may be suggested. 

1. The vegetation that recolonizes the land will in all possibility 
be different than that which previously grew in the area. Most likely 
many previous floral components would now find the area inhospitable. 

2. Recolonization of the habitat by organisms would theoretically 


(2):i09-ii3, 196 } 


SAND CREEK ARGYNNIS 


113 


be possible from any portion of the surrounding regions. It may be 
expressed by saying that recoionization could be expected to be muUi- 
dkectional unless some factor prevented recolonization from one or 
more directions. 

3 . Such recently recolonized areas would contain mixed populations 
the components of which may have moved in from any of the sur- 
rounding regions. 

4. If such heterogeneous populations are studied before adequate 
time for stabilization has passed, it is predicted that great variation in 
individual phenotype will be present. 

Expressing this in terms more directly related to the Argynnis 
populations of the Sand Creek Basin, it is tentatively concluded that 
the variability of these several populations is in part a result of the 
recency with which they have established themselves in this area follow- 
ing a major change in habitat, and in part a result of recolonization by 
an unknown number of varying populations from the surrounding 
regions. 

Less tangible are two other possibilities. One is that the time during 
which the Sand Creek Basin was unsuitable for habitation by Argynnis 
was of sufficient duration to allow the surrounding populations which 
it separated, to diverge to some, extent. The other is that the recent 
forest cover of the pumice soils may differ sufficiently in its selective 
factors io that the eventual stable population may when developed 
differ from those around it. 

Thanks are due to the National Park Service, and in particular to 
Park Naturalist Bruce Black, for extensive courtesies and helpful 
information. 


REFERENCES 

BRIGGS, LYMAN J. 1962., When Mt. Mazama lost its top. National 
Geographic, 122 (1): 128-133. 

MOECK, ARTHUR H. 1957. Geographic variability in Speyeria. The 
Milwaukee Entomological Society, 48 pp. 

WILLIAMS, HOWELL. 1961. Crater Lake — The story of its origin. 
University of California Press, Berkeley & Los Angeles. XII + 98 pp., 
10 pits., 8 figs. 


Journal of Research on the Lepidoptera 1(2) 1963 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 7563 


CATERPILLAR VERSUS DINOSAUR? 

THEODORE H. EATON, JR. 

Museum of Natural History and Department 
of Zoology, University of Kansas 

A PAPER BY S. E. Flanders in the first issue of The Journal of 
Research on the Lepidoptera (Flanders, 1962) bears the intriguing 
title, "Did the caterpillar exterminate the giant reptile?” In his dis- 
cussion, the "abrupt end of the Age of Reptiles during the Cretaceous 
period is ascribed to a newly emerged order of insects, the Lepidoptera, 
on the supposition that for a brief period it regulated the world’s 
supply of plant life at starvation levels for the dependent reptiles.” 
The argument for this hypothesis rests in part on modern examples 
of large-scale devastation of certain species of plants by the larvae of 
Lepidoptera, and in part on assumptions concerning the time of origin 
and expansion of this order, the nature of plant life during the Cretaceous 
period, and the habits and relationships of the "giant reptile.” 

Many explanations have been proposed for the extinction of dino- 
saurs and several other groups of reptiles at the end of the Cretaceous, 
but there is not yet any general agreement as to the most important 
causes. The proposals are conjectures, having little or no direct evidence 
to support them. The theory under discussion adds another to the list. 
The consensus among paleontologists now is that one or two "causes” 
are not sufficient, and that the extinction of these animals is a more 
complicated phenomenon than it has seemed. While it is possible that 
destruction of food-plants by caterpillars has at times contributed to 
extinction of some herbivorous reptiles, and thus indirectly to that of 
their predators, the present writer thinks that a close look at what 
happened in the Mesozoic might be helpful in judging this hypothesis. 

Our knowledge of fossil Lepidoptera is' scanty, but the earliest 
known moth appears to be Loses triassica, described from Upper Triassic 
beds in Queensland (Tindale, 1945). It is known only from a fore and 
hind wing, of which the venation agrees broadly with that of Jugatae 
(Homoneura); Tindale placed it in a new, more primitive suborder, 
Eoneura. Thus the order must have originated more than a hundred 
million years before the extinction of dinosaurs, although leaf-eating 
caterpillars might not have characterized the earliest genera and families 
of moths. Reports of moths and butterflies from the Jurassic litho- 
graphic limestone in Bavaria are based on specimens now attributed 


114 


i(2):ii4-ii 6, ip6j 


CATERPILLAR VS. DINOSAUR? 


115 


to other orders, especially Homoptera. Unless more information appears 
in Frank M. Carpenter s forthcoming volume on fossil insects in the 
"Treatise on Invertebrate Paleontology," we can say nothing specific 
about Lepidoptera of the Jurassic and Cretaceous; none are mentioned 
in the "Traite de Paleontologie,” vol. Ill (1953). In the Cenozoic, 
however, moths and butterflies of modern families are known, apparently 
from the Eocene on. 

It is reasonable to suppose that the rise of Lepidoptera coincided 
with that of flowering plants. These plants are known first from the 
Jurassic, but at least sixteen modern families are represented in Lower 
Cretaceous rocks; by the late Cretaceous many existing species of trees 
had appeared. The length of the Cretaceous was about 70 million years. 
From its close to the present time was a little less, say 63 million years, 
according to recent determinations (Kulp, 1961). But we cannot, in 
the nature of the case, show evidence that caterpillars appeared in great 
numbers in the late Cretaceous, or that they were then so free of para- 
sites or predators that they could, by worldwide devastation of all sorts 
of plants, bring starvation upon the reptiles. 

Some of the statements of Flanders concerning reptiles bear comment 
from the viewpoint of a paleontologist. For instance, "The inherent 
weakness of the reptile was an extraordinary need for an abundance of 
plant material . . . The small size of today’s descendent reptiles, the 
vegetarian turtle, the predatory crocodile, the snake, and the lizard, is 
evidence 'of the giant reptile’s elimination by starvation and predation.’’ 
A sauropod, hadrosaur, ankylosaur or ceratopsian dinosaur may have 
eaten a large quantity of vegetation, the amount depending on size and 
activity. These animals were present in the late Cretaceous and occupied 
a position comparable in some ways to that of the browsing mammals 
of the Cenozoic. But their extinction does not seem to have been abrupt; 
it was a slow, uneven decline probably extending through millions of 
years, and does not differ so far as we know from the decline and 
disappearance of various other groups of animals at earlier and later 
times. The flying pterosaurs, whose food was probably fish, declined 
and died out at about the same time as the dinosaurs. So also did 
mosasaurs, a specialized line of giant marine fish-eating lizards; so too 
the very different aquatic plesiosaurs, as well as a few late Cretaceous 
ichthyosaurs and carnivorous dinosaurs. 

No living reptiles are directly descended from any of the above- 
mentioned Mesozoic reptiles. Turtles and crocodiles originated in the 
Triassic and have continued with no fundamental changes to the present 
time. Lizards are known first from the Jurassic, snakes from the Creta- 
ceous, and their size cannot be taken as evidence of any changes among 
the giant reptiles. These giant reptiles were clearly not a single natural 
group, nor, on the other hand, were all members of the orders to which 
they belonged of great size. In the two orders of dinosaurs, for example, 
there were some species, both early and late, that failed to reach the 
size of an average alligator. 


116 


EATON 


/. Res. Leptd. 


Efforts to explain the extinction of dinosaurs and various other 
reptiles in the Cretaceous period ought to take into account a number 
of possible causes. The one proposed by Flanders does not give the 
answer to several problems, such as the disappearance of various un- 
related and ecologically different groups, not necessarily of large 
animals, and the survival beyond the Cretaceous of certain other 
reptiles, not necessarily small. There is no evidence of either a biological 
or a geological catastrophe of large proportions at the close of the 
Mesozoic, unless the slow retreat of epicontinental seas in some areas 
can be so described. Probably the answer will eventually be found in a 
combination of many factors. 


LITERATURE CITED 

FLANDERS, S. E. 1962. Did the caterpillar exterminate the giant reptile? 
J. Res. Lepid., 1 (1): 85-88. 

KULP, J. LAURENCE. 1961. Geologic time scale. Science, 133 (3459) : 
1105-1114. 

TINDALE, N. B. 1945* Triassic inseas of Queensland. Rroc. Roy. Soc. 
Queensland, 56 (5): 37-46. 


Journal of Research on the Lepidoptera 1(2) :117-123, 1963 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 196} 


GEOGRAPHICAL DISTRIBUTION AND VARIATION 
OF THE GENUS ARGYNNIS 

I INTRODUCTION 
IL ARGYNNIS IDALIA 
WILLIAM HOVANITZ 

California Arboretum Foundation, Inc., Arcadia, California, and 
Los Angeles, State College, Los Angeles, California 


L INTRODUCTION 

The Genus Argynnis in the comprehensive sense covers a wide 

variety of form and size in Nymphalid butterflies. The geographical 
distribution of the genus as a whole is somewhat comparable to that 
of Colias but is actually wider in some areas due to a tropical group 
in the eastern hemisphere. A natural desire to have a greater similarity 
of type within a "genus” and a desire of some workers to "split” to the 
ultimate has left the genus in a diverse state of fragmentation. This has 
been aided by regional preferences and the difficulty for most Lepid- 
opterists to observe and study members of the group from continents 
other than their own. The members of the genus from the smaller 
isolated continents such as North America and South America have 
apparently either diverged slightly from the type on the largest land 
mass (Eurasia) or they have retained more primitive features. In 
North America only two of the several subdivisions of type within the 
Argynnis appear ever to have arrived for colonization. These two have 
diverged to such an extent on that continent that residents of that 
continent of natural inclination should consider them as of different 
genera {Boloria and Speyeria) . One of these {Speyeria) is only slightly 
different from one of the species groups of Eurasian Argynnis, namely, 
aglaja, adippe, niohe, etc. and placed by some in a genus Brenthis which 
is not the usage of Brenthis used in North America by others. 

The variety of the genus in Eurasia is much greater than in North 
America. It is natural to expect that the types present in Eurasia which 
would be most like those now present in North America would be 
those whose climatic adaptations would be the most similar to the 
conditions present in the area where a land bridge exists now or did 
exist in the immediate past. This is the area of Alaska and adjacent 
Siberia. The Boloria group is the best adapted to the climates of this 
area and may even now be considered for all practical purposes to have 
"continuity” across the barrier at the Bering straits. With the possi- 
bility of gene flow not very remote, it is not difficult to see why Boloria 


117 


118 


HOVANITZ 


/. Res. Lepid. 


of North America and Asia (Europe) should be much alike. In fact, 
several species of Boloria can and do go under the same species name 
on both continents, just as do three species of Colias which have the 
same geographical relationships to the land bridge. 

The species which are the most remote in their climatic preferences 
from the climates of the Bering area are those species which are the most 
divergent on the two continental areas; these are the most southern 
types. Argynnis paphia is the member of a southern subgenus of 
Argynnis in Eurasia and this subgenus apparently never colonized 
America. Instead, a type similar to aglaja apparently did colonize 
America and from this presumably all the American Argynnids of 
the larger {Speyeria) type subsequently were derived. It is an hypo- 
thesis that the most extreme types in North America, such as idalia, 
diana and nokomts, which bear considerable pattern and color relation- 
ship to Eurasian species such as paphia, sagana, childreni, etc. were 
derived independently from them from aglaja — like ancestors under the 
selective influence of the comparable conditions of climate to which 
they were subjected (hot-humid summers). Comparative correlations 
of the facies of the butterflies with climates at different parts of the 
world will be made later in this series. 

The Argynnis of South America bear no close relationship apparent- 
ly to the Argynnis of any other part of the world. There is no easy way 
at the present rime for them to have arrived there by migration from 
other continents. Only two alternate proposals seem possible. The first 
of these is that these Argynnis were isolated many years ago (the early 
Tertiary) from the other Argynnis of the world and that they arrived 
there by migration from North America at a time when North America 
was inhabited by a more primitive type. They might also have arrived 
from Africa early in the evolution of the genus should there be reason 
to believe seriously in the past movement of South America away from 
a more close proximity to Africa. The second proposal on the origin 
of the South American Argynnis is one of independent origin in South 
America from some tropical or subtropical relative of the North 
American Argynnis. This proposal seems very weak in view of the lack 
of any close relative that would seem to be a likely ancestor. Dryas 
(Colaenis) , Dione and Euptoieta seem to be out of the question, though 
this may be only based upon superficial appearances. They feed upon 
similar plants, a factor greatly favoring the argument, especially with 
regard to Euptoieta which inhabits cold country in the Andes, feeds on 
violets and has much the same habits as Argynnis itself. 

The question comes up as to what is a genus. Should the entire 
group which has been pictured here be considered one genus, or should 
it be raised up to a sub-family or even a family. If the latter, then 
should the major groups be raised from subgeneric level to the generic.^ 
If so, then how many groups at this level should be recognized? The 
question is then condensed to one problem. There really is no biolo- 
gical entity which can be known as a genus. All hopes to the contrary, 


(2);ii7-I23, 1963 


ARGYNNIS DISTRIBUTION 


119 


the genus is solely a matter of convenience in nomenclature, which 
should show as much as possible, phylogenetic relationships. The 
system should start at the bottom of the classification heirarchy. If it is 
known what should be considered species, sufficient related species 
ought to be put into a genus to make a reasonable group on the basis 
of numbers and morphological (genetical) similarity. Should too few 
species be put into a genus the advantages of the binomial nomen- 
clature are destroyed, in much the same way as would occur if every 
person had the family name Smith. A further criterion, in addition to 
size and uniformity, is geographical coverage. It is a natural phenome- 
non that isolation leads to differentiation. Related members of a species 
group may diverge slightly due to continental isolation and yet in all 
other ways be closely knit as a single group. The decision here must 
rest on the needs of the taxonomist in showing relationships and the 
usefulness to biologists. Nomenclature bears one major value: that of 
usefulness. If different generic names were to be used on each con- 
tinent, a degree of provincialism would develop which would be hard 
to penetrate; local butterfly books in North America and in Europe 
have now diverged to such an extent in terminology that the only way 
a novice can determine relationships between the butterflies of Europe 
and North America is to look at the specimens — -an ironic turn of 
events since that should be the purpose of nomenclature. As a result 
therefore, it is the hope that there may be a return to the larger genera 
of the past; this is the thesis upon which the use of Argynnis here is 
based, without necessarily accepting the assumption that the generic 
limits and the characters used for their delimitation should remain as 
in the past if more specific future work should show the need for 
change. 

IE ARGYNNIS (SPEYERIA) ID ALIA 
This representative of the North American Argynnis differs in 
appearance from the usual Speyeria probably more than any other 
except possibly for A. diana. The details of its pattern which are dis- 
tinctive and different from the standard Argynnis pattern are the greatly 
increased black pigment around the borders of the wings, even extending 
to the basal section and nearly covering the hind wings, but at the same 
time leaving free of black pigmentation, the marginal and submarginal 
rows of spots ( fig. 1 and 2 ) . The general facies therefore is distinctive and 
different from any other Argynnis. At the same time, the black pattern 
elements of the center of the fore wing are reduced rather than ex- 
panded as in all other related Argynnis. It therefore shows a develop- 
ment of a pattern not only unlike that of the Argynnis, including the 
subgenus Speyeria as indicated by Dos Passos and Grey, but a trend 
in a diametrically opposed direction. The distinctiveness of this species 
is such as to need no real description or detailed analysis. Likewise, from 
the study of specimens throughout the geographic range of the species, 
there are no geographical variations apparent, though there are in- 


120 


HOVANITZ 


/. Re$. Lepid. 



Fig. 1. Argynnis idalia, upper side; top: male; bottom: female, from Oak 
Park, Illinois, July 8, 1906. F. S. Daggett. 


i(2):ii7-i2h 19^3 


ARGYNNIS DISTRIBUTION 


121 



ilfjflllliilililigF 




iisSllfiL 


iSisiSssSssilism 


immmmmmmmmmmm 

|bs::ss;s::l 

^^asms: 

g^^SSSSSSSi 
rfw slow******* 

W ^Ms^sssi 


Fig. 2. Argynnis idalia, same as fig. 1 only lower side. 


122 


HOVANITZ 


/. Res, Lepid, 



SCALE 


1000 FOOT CONTOUR 


Fig. 3. Map of North America showing the distribution of Argynnis idalia. 


(2):ii7-i2}, 1963 


ARGYNNIS DISTRIBUTION 


123 


dividual differences in size and slight differences in the pattern observed 
on an individual basis. Perhaps statistical differences could, and will, 
be detected in the future. 

The distribution of this species is limited to a central and eastern 
location across the United States and a small part of Canada. The 
biological reasons for this restricted distributional range are not known. 
It can be seen from the map (fig. 3) that the species ranges from New 
England (specimens which I have seen come from as far as Massa- 
chusetts though records indicate Portland, Maine as the furthest point 
reached north and east), southward to North Carolina (probably erro- 
neous records indicate Georgia), and west to Colorado and North 
Dakota (probably erroneous records indicate Montana). There are 
records or indications in old literature for Arkansas but I have seen 
no specimens from there. The species seem to prefer the area south of 
the Canadian life zone, that is, the Transition life zone of sorts, though 
it is pretty hard to outline any definite climatic zonation. Records 
indicate that it is to be found in most of the southern counties of 
Michigan and the southern counties of Minnesota. Scudder (1889) 
writes: "This butterfly belongs to the Alleghanian fauna, though its 
distribution appears to be somewhat irregular. It inhabits lowlands and 
is much more abundant in the extreme eastern portion of its range than 
elsewhere, unless it be the western prairies.” Mention of it in Louisiana 
by Strecker is most probably in error. 

A further discussion of the range of this species will be deferred 
until the maps of other species have been published, which will be 
used for comparison. 


REFERENCES 

DOS PASSOS, C. F. AND L. P. GREY. 1947. Systematic catalogue of 
Speyeria (Lepidoptera, Nymphalidae) with designations of types and 
fixations of type localities. Amer. Mus. Novitiates No. 1370. 

— — — — 1945. A genitalic survey of Argynninae (Lepidoptera, Nymph- 

alidae). Amer. Mus. Novitates No. 1296. 

MACY, R. W. AND H. H. SHEPARD. 1941. Butterflies (of Minnesota). 
Univ. Minn., Minneapolis. 

MOORE, SHERMAN. I960. A revised annotated list of the butterflies of 
Michigan. Occasional Papers of the Museum of Zoology, University of 
Michigan. No. 617. 

SCUDDER, S. 1889. The Butterflies of the Eastern United States. Cambridge, 

Mass. 


Journal of Research on the Lepidoptera 


1 ( 2 ) : 124 - 134 , 1963 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
© Copyright 196} 


THE RELATION OF PIERIS VIRGINIENSIS EDW. 
TO PIERIS NAPI L. 

SPECIES FORMATION IN PIERIS? 

WILLIAM HOVANITZ 

California Arboretum Foundation, Inc., Arcadia, California, and 
Los Angeles State College, Los Angeles, California 

In a previous paper (Hovanitz, 1962), it was indicated that 
a fuller discussion of the relationship of Pieris napi L, to Pieris virgin- 
iensis Edw. would be forthcoming. In that paper, the map showing the 
distribution of P. napi included the distribution of P. virginiensis but 
a closer distinction between the two was not indicated because of the 
reduced size of the map included. A more detailed map showing the 
relationships between the two biological entities under discussion is 
included with this paper, together with a description of the known 
facts of the lines of separation of the two "species” or races, and a 
summary of the events that have most probably occurred in relation 
to the two groups up to the present time. 

On the map in the previous paper (Hovanitz 1962), the range 
of Pieris napi L. for the eastern United States included virginiensis as 
well as napi itself. The race of napi with which we are concerned here 
is known as oleracea. This race blends northwards and westwards into 
other races by imperceptible gradations. Pieris napi oleracea (known 
also by the summer brood names, aestiva Edw., hyemalis Edw. and 
acadica Edw.) may be considered the name for the race of napi east 
of 90° Longtitude and south of 50° N. latitude, more or less, with the 
exception of the area south of New York state (38° N). Other names 
can be found in McHenry (1962). The area south of New York 
state and east of 90° Longitude is occupied by the race or species 
virginiensis. In New York state and other points in the vicinity, or close 
to the southern limits of oleracea, there are populations which in some 
cases appear to be more closely related to virginiensis than to oleracea, 
and in other cases, are more closely related to oleracea with occasional 
individuals similar to virginiensis. The status of the line of contact 
between oleracea and virginiensis is not one of free and even blending 
such as might be expected of freely interbreeding individuals in a case 
of solely geographical isolation. Populations are rather clearly either 
oleracea or virginiensis as the case may be, with only occasional examples 
of intermediates giving some hint of the nature of introgression at the 
line of contact. 


124 


( 2 ) -. 124 - 1 34 . 19^3 


PIERIS NAPI RACES 


125 


The contact of these two species or races bears a close relationship 
to the similar situation between the species or races Basilarchia astyanax 
and B. arthemis in the same general area (Hovanitz 1949). 

Specimens of P. napi studied in various collections have permitted 
a map to be drawn showing the detailed distribution of oleracea and 
virginiensis (fig. 1). On this map, locations where oleracea (or napi in 
the species sense) are known are shown by the letter N and virginiensis 
locations are shown by the letter V. In addition to the indications on 
the map, there are reasons for assuming that each of these races overlaps 
the other even greater than seems indicated on the map. Due to lack of 
specific data (e.g. "New York”), some of these indications could not 
be shown on the map. These details will be mentioned below. 

On the map, it can be seen that oleracea extends southwards in the 
mountains of New England to Connecticut where it comes in contact 
with virginiensis or something like virginiensis . It is doubtful that these 
specimens are truly virginiensis, though they do bear some resemblance 
to them. Instead they are more likely abberant oleracea, which are 
genetically tending in the direction of virginiensis through introgression. 
In addition, the habitat change out of the Canadian forest typical of 
oleracea to the "Transition” lowlands might lead to a phenotypic 
alteration in facies. Populations of considerable size are definitely known 
at the present time in Vermont and New Hampshire. They are not now 
known in the vicinity of Boston, Mass, though they were common there 
before 1850. The following bears witness to a reduction in distribution 
of napi in the 1800 s: 

From Scudder (1889) : 

"This butterfly is a member of the Canadian fauna . . 

"It is found throughout New England although seldom abundant south 
of the annual isotherm of 48°. Northward and eastward it is everywhere abund- 
ant and it continues to be so as far south as Williamstown, Mass. (Scudder), 
Dublin, N. H. (Leonard) and Portland, Me. (Scudder, Verrill). South of 42° 
N. L. or the northern boundary of Connecticut, it rarely occurs, although it 
has been taken at Farmington, Conn. (Norton), Newport, R. I. (Miss 
Coggeshall teste Higginson) and Fire Island Beach, Long Island (Smith). 
Even in the north, however, there seems to be some local causes for its 
abundance.” 

"Mr. Lintner, writing in 1864, says that in Schoharie, N.Y., 'it was 
extremely rare until recently,’ and previous to 1857, 'he had taken it but 
once in three years . . "Mr. Bacon of Natick, Mass, says that the insect by 
no means disturbs cabbages and turnips as it did fifteen or twenty years ago.” 

"I recollect once seeing the college yard in Cambridge, I think it was 
about 1857, fairly swarming with P. oleracea. It is now never found, to my 
knowledge, anywhere in the region about Boston, and I think it is wholly 
confined to the less cultivated and especially the hilly distrias of New England.” 

FromKlots (1951): 

"Napi has . . . suffered a great restriaion in habitat, and probably in 
habits, for it is now almost entirely limited to shaded Canadian Zone forest 
. . "Not recorded south of the Catskill Mountains in New York”. 


126 


HOVANITZ 


/. Res. Lepid. 


From Klots (1935) of Edwards, 

"He mentions a female taken by Mead in the last week of June at Stony 
Clove in the Catskill Mountains . . . many eggs were obtained . . . from the 
chrysalis an oleracea emerged.” "Stony Clove is in excellent territory for napi 
{oleracea) but I know of no records of virginiensis from there.” "I have . . . 
seen specimens of virginiensis from Big Indian Valley in the Catskills, where 
the environment is more suited to virginiensis.” 

"In 1931, near McLean, Tompkins County, New York, I was able to 
obtain undoubtedly authentic eggs, larvae and chrysalis of virginiensis, through 
watching females ovipositing on Dentaria diphylla Michx.” 

From Fiske ( 1901 ) : 

"Pieris napi. once common enough to be injurious to cabbages and other 
cruciferous plants throughout New England, has since the introduction of rapae 
become one of our rarest butterflies. Its range is now restricted to the northern 
and mountainous portions, and even in the fortresses of the White Mountains it 
is a scarce insea.” 

The eighteenth century idea of a "species” contributed a lot to a 
general confusion of the nature of the relationship between oleracea and 
virginiensis. It was then, and too often now, the general idea that a 
species was represented by a specific morphological type, without regard 
to the variation which a population may have within itself. Some of 
the best collectors and breeders of Lepidoptera have therefore become 
confused, having the all pervading desire to designate some biological 
unit as either a "species”, subspecies or variety when the true relation- 
ship may not be possible with only these tools of nomenclature. For 
example, Edwards (1881) says: 

"Virginiensis . . . has become a true species, although unquestionably, in 
a higher latitude, it appears as an occasional aberration only of Oleracea.” 

Also, Klots (1951) says: 

"Until very recently, virginiensis has been confused with napi, but it is 
now known to be a distinct species of more southern (Transition Zone) 
distribution.” 

Edwards (1868-1872) says: 

"In the Kanawha district, it replaces Oleracea which is yet unknown there. 
It is not uncommon in the month of May, frequenting open woods rather than 
gardens, and in this respect differing in habit from the allied species. I have 
never met with it later than June, though Oleracea, in the Northern States is 
most abundant after that month and continues breeding till the early autumn 
frosts.” "I have received specimens of Virginiensis from Mr. Wm. Saunders of 
London, Canada and am informed by him that it is there a rare insect.” 

DIFFERENCES BETWEEN P. NAPI OLERACEA AND 
P. NAPI VIRGINIENSIS 

The primary visible differences between P. napi oleracea and P. 
napi virginiensis can be observed in figs. 2 and 3. In figure 2 is shown 
the male and female of the upper side of virginiensis to the left; at the 
same location in figure 3 is shown the under side. These specimens are 


(2):i24-n4, 196} 


PIERIS NAPI RACES 


127 



Fig. 1. Map showing the distribution in the eastern part of North America 
of the introgression area of Pieris napi races virginiensis and oleracea. 


from near Pittsburgh, which although toward the north end of the 
range of virginiensis are quite typical of the populations south through 
the Appalachian mountains. The pattern of the upper side female 
should be noted as quite typical of this race, and which is only rarely 
present in oleracea. Of critical distinction, the pattern of the under side 
of the wings should be noted. The veins are outlined in a tan pigmen- 
tation, not heavy but rather broad in expanse, in appearance unlike 
any other race of napi. 

P. napi oleracea is shown in figures 2 and 3 by the central two 
specimens; these are from southern New Hampshire which is near the 


128 


HOVANITZ 



J. Res. Lepid. 


« 2 
§ a 
6 w 
^ On 

..U 


o § 

a M 
.. B 

Ph S 
H • 


m 

0\ 

X 

§ 

^ a 


^1 S 


,3 3 


V N 

S m 

^ G\ 
G ^ 
P rn 


a ph‘ 

O G\ 

H 


1 ^ 

dl ^ 

e§ 

^ e« 
^ « 
5 W 

-4 

0 aJ 

t- a 

1 ^ 


fN 

E 


(2); i 24-134> 19^3 


PIERIS NAPI RACES 


129 



Same as fig. 1, but all lower side fades' 


130 


HOVANITZ 


/. Res. Lepid. 


southern part of the distributional range of this species. They are how- 
ever typical in appearance. Note the difference between the two races 
on both the upper and lower surfaces: virginiensis is more of chalky 
white on the upper side as compared with oleracea; the wings on the 
upper side tend to be veined with black near the borders and cell in 
oleracea which is replaced with a brownish ''rapae-ty^t' pattern in 
virginiensis; on the under side the veins are heavily lined with black in 
oleracea in a way distinctive from that of virginiensis; and there is a 
yellowish suffusion on the under side of the wings in oleracea not 
present in virginiensis. On the other hand, the appearance of oleracea 
changes during the seasons while that of virginiensis does not. There is 
no indication that virginiensis has more than the one generation per 
year, this occurring in early May whereas oleracea has a succession of 
generations starting in April and extending throughout the year until 
the cold weather of the autumn. During this extended period the weather 
conditions are quite different, and the effects of this on the appearance 
of oleracea are profound. The appearance of the adults emerged during 
the warmer part of the season are quite different. A specimen of the 
summer brood is shown in figures 2 and 3 at the upper right. Note the 
slightly increased size but more important the almost complete absence 
of any pigment other than white on the wing surfaces. These summer 
forms have been given the names hyemalis, acadica and aestiva by 
Edwards for the general area under consideration. As a racial name, the 
only one of any significance however is oleracea. Throughout the ranges 
of these races, there is fairly good uniformity in pattern and character- 
istics; however, it should be pointed out that within each of these races 
and especially oleracea there is a great amount of variation which tends 
to blend the two together. For example, a specimen at the bottom right 
in figures 2 and 3 is from a population of oleracea but shows the "rapae- 
type” pattern of virginiensis. The underside of the wings however makes 
clear that the specimen is oleracea. 

Areas of typical oleracea may be described as follows: The Can- 
adian forest areas ranging from northern Michigan across northern 
Ontario and Quebec into eastern Quebec, New Brunswick, Nova Scotia, 
Maine and the mountainous parts of the New England states as far as 
Massachusetts. South of these points and until the typical populations of 
virginiensis are reached in New York state, Ohio and Pennsylvania^ the 
populations apparently are in a state of uncertainty regarding a complete 
divergence from the virginiensis or the oleracea genome and all the 
things pertaining thereto, including in addition to pattern differences, 
the physiological alterations required for the more southern existence, 
the univoltine life cycle and the changed food plant preferences. These 
intermediate populations lie in a line ranging from Michigan, across 
southern Ontario to Montreal in Quebec and southward to the Catskill 
Mountains of New York. Some of these can be distinguished as oleracea 
with virginiensis tendencies and others as virginiensis with oleracea 
tendencies, though the former outrank the latter. Populations which 


(2); i 24-134, 1963 


PIERIS NAPI RACES 


131 


would be considered oleracea with virginiensis tendencies would be the 
following, with an indication of the extent of the tendency: 

Michigan: Emmett Co., north part of southern peninsula (slight) 

Ontario: region around London, Huntsville, Sydney Field station, 
Marmora, Trenton ( considerable ) 

— - — : Aylmer, Bells Corners, Brittania, South March, Merivale, 

Constance Bay, Ottawa West (slight) 

Quebec: Harrington Lake, Gatineau Park (slight) 

Massachusetts: Cummington, Whately, Franklin Co., Mt. Greylock 

Populations which would be considered virginiensis with oleracea 
tendencies would be: 

Quebec: He Perrot (Montreal) 

New York: the Adirondacks, the Catskills, Ilion, McLean 

Ontario: Hamilton 

Populations with little sign of intergradation on the northern end 
of the range of virginiensis'. 

Pennsylvania: Scranton, New Brighton, Pittsburgh, Forbes Forest, 
Charlevoi, Showville, Washington Co., Fayette Co., Lawrence Co. 

Ohio: Canton 

West Virginia: Forks of the Kanawha, Coalburgh 
HABITS OF THE RACES 

The 'rather distinct break between the geographical ranges of 
oleracea and virginiensis would seem to need some isolation barrier for 
the maintenance of its integrity. This barrier is clearly not one of water 
or mountain but rather one of habitat selection and the accompanying 
biological changes in the physiology of the races concerned. For some 
idea of the field habits of these races, we may refer to other authors: 

Klots ( 1951 ) says of virginiensis'. 

"Until very recently, virginiensis has been confused with napi, but it is 
now known to be a distinct species of more southern (Transition Zone) 
distribution. Its dark markings are a more smoky, diffuse brown and it lacks 
the yellowish tint beneath, on the HW and apex of the FW, which napi 
usually has. It is single brooded. More even than napi, it is limited to woods 
and very local.” 

"Food: — Toothwort {Dent aria diphylla) , probably other Dentaria. One 
brood. Adults in early May (central New York.” 

"Range:— Transition Zone, Ontario, central New England and New York, 
s. to Virginia (TL. Kanawha, West Virginia) 

He says of napi oleracea'. 

'^Napi has . . . suffered a great restriction in habitat, and probably in 
habits, for it is now almost entirely limited to shaded Canadian Zone forest 
... A virginiensis lacks yellowish beneath on the HW and apex of the FW 
and has the scaling along the veins of the HW and costa and apex of FW 
beneath much more diffuse and powdery looking. A rare dark form of P. rapae 
lacks dark spots above, but never has the dark-lined veins of napi, and has 
heavier scaled, less fragile looking wings.” 

Food. Formerly, at least, nearly any cultivated or wild cruciferae; now 
probably chiefly limited to native, woodland species, such as toothworts 
(Dentaria), Rock cresses (Arabis), Winter and Water cresses (Barbarea), etc. 


132 


HOVANITZ 


J. Res. Lepid. 


Three broods. Adults in late April (Massachusetts).” "Not recorded s. of 
the Catskill Mountains in New York.” 

Clark (1951) says of virginiensis: 

"Range :---One record, Frederick County northwest of Cross Junaion on 
the Bloomery Road (Rank 698) about 200 feet east of the West Virginia 
line, April 24, 1938.” 

"Variation — This species varies considerably in the intensity of the 
bordering of the veins on the under side of the hind wings. This bordering is 
pale in ail the individuals from Virginia and adjacent West Virginia that we 
have seen, some having the under side of the hind wings almost immaculate 
white.” 

"Occurrence: — Confined to rich deciduous woods and extremely local, 
occurring in limited numbers at usually widely separated localities. The reduction 
in the numbers of Pieris virginiensis and its present occurrence only in widely 
separated localities are undoubtedly the result of deforestation, which has 
greatly reduced the areas in which it is possible for it to maintain itself.” 
"Pieris virginiensis keeps strictly to the woods.” 

"Season: — One brood. This species appears shortly after the middle of 
April and flies until about the middle of May.” 

Clench (1958) writes of the same: 

"Pieris virginiensis . . . common in woods . . . one brood: Late April to 
mid-May. One . . . capture ... on 4 June, 1958 is surely a freak occurrence . .” 

Scudder ( 1889) writes of the habits of oleracea: 

"Food plants. This caterpillar feeds on various cultivated cruciferous plants, 
such as turnip {Brassica rapa) , cabbage {B. oleracea), radish {Raphanus 
sativa) , horseradish {Nasturtium armoracia) and mustard {S inapis ) . Mr. 
Lintner considers that it prefers turnip to cabbage, for he once obtained fifty 
eggs fom a turnip bed and could find none on adjacent cabbage plants of the 
same age, and this has been my experience.” Hoy, on the other hand, thinks 
it is found mostly on mustard. But it also occurs on some of our native plants, 
such as Arabis drummondi (Couper) and A. perfoliata (Fletcher), and Fitch 
says it occurs abundantly on the water cress {Barbarea vulgaris)." 

"Life History. This butterfly, which appears to be generally triple brooded, 
passes the winter in the chrysalis, the first brood has been seen as early as 
April 18, but usually appears between April 27 and May 9, a week or there- 
abouts after P. rapae. For several years Fitch observed it at East Greenwich, 
N. Y. for the first time on May 8 and 9 and only once as early as May 6; in 
northern localities it is sometimes as late as the third week in May. It usually 
becomes abundant between the 10th and 15th of the month and continues 
unti the end of the first week in June. The eggs are laid during the last half 
of May and early in June, and hatch in from 5 to 10 days . .. . the second 
brood . . . makes its advent during the last days of June or early July . . ., this 
brood in oleracea seems never to be very abundant . . . and to maintain itself 
for a comparatively short time; it becomes common by the end of the first 
week in July and sometimes disappears . . . before the end of the month . . . 
"The third brood appears as early as the last days of July but usually not before 
August; it . . . lasts until early September . . . Occasional specimens, . . . 
disclose butterflies late in September or early in Oaober.” "The species must 
be triple brooded in the north.” 

Edwards (1868-1872) wmts oi virginiensis: 

"In the Kanawha district it replaces Oleracea which is yet unknown 
there. It is not uncommon in the month of May, frequenting open woods 
rather than gardens, and in this respect differing in habit from the allied 
species. I have never met with it later than June, though Oleracea in the 


(2):i24-i34, 196} 


PIERIS NAPI RACES 


133 


Northern states is most abundant after that month and continues breeding 
til the early autumn frosts.” "I have received specimens of Virginiensis from 
Mr. Wm. Saunders of London, Canada and am informed by him that it is 
there a rare insect.” 

These data would tend to confirm the evidence derived from the 
data of museum specimens that oleracea is indeed a muitivoltine race 
which breeds from April all through the year until the Autumn frosts. 
Our own personal experience on napi in California does not conflict 
with this opinion. Here napi {venosa and castoria) are continuously 
brooded when possible in relation to temperature and moisture condi- 
tions. They go into diapause when conditions of the food plant are un- 
favorable; this is usually in the summer when the food plant {Dentaria) 
dries up. In fact much of the first winter generation does likewise as 
Dentaria does not stay green long. This accounts for the summer or late 
spring generations being less abundant than the winter or early spring 
generations. A similar observation has been made of the generations of 
napi in New England. 

The food plant difference between the races oleracea and virginiensis 
is important in only one way, namely in the restriction of habitat of 
napi to a different ecological niche. Dentaria is a plant of the open 
woods and partial shade. The flight period of virginiensis must be 
adjusted to meet the proper conditions of moisture requirement for the 
plant as well as for the degree of light availability in the deciduous 
forest where early spring may allow more light in the forest than in 
summer. The restriction of virginiensis to the "Transition” life zone 
forest may be correlated with this light and temperature relationship. 
The northern forest is cooler all summer long and the temperature needs 
of oleracea can be met usually in the open areas as well as in the partial 
shade of the forest. This would account for the different flight habits 
of the two races. In California, napi ( venosa and castoria ) fly only in the 
cool canyons where there is partial shade even though there is abundant 
food available for them outside in the form of wild mustards. Instead 
they are restricted to Dentaria of the shaded woods, while Pieris rapae 
and P. protodice fly only a few yards away in the direct sunlight and 
feed upon mustards. 

The food plant of oleracea so often indicated in the literature is 
winter or water cress {Barb area) which is apparently restricted to the 
vicinity of strearqs. There is indication that in the Catskill Mountains 
(New York) there may be a population of virginiensis on Dentaria 
while sympatrically there are populations of oleracea on Barbarea. If 
this is so, an excellent study could be developed to show the nature of the 
racial or specific isolation of these two at a region where both maintain 
reasonably distinct populations in close contact. The fact that oleracea 
flies in April, virginiensis in May and oleracea again in June and July 
indicates another additive isolation factor which would help to maintain 
the species or racial distinctness. 

By way of conclusion, the data presented show rather conclusively 
that Pieris napi oleracea and Pieris napi virginiensis are not as com- 


134 


HOVANITZ 


/, Res. Lepid, 


pletely separated as one might wish "species” to be but that they are 
not either fully racial or subspecific in the sense that their populations 
blend into one another as by a continual gene flow. It is therefore a 
matter of personal opinion whether they should be considered species or 
races for nomenclatorial purposes. For these purposes it is impossible 
to stand on the middle ground as a choice must be made. I choose to 
use the racial designation for the reason that the groups are almost 
completely allopatric, that is, the populations are geographically separ- 
ated and do not occur in the same place with certainty without inter- 
breeding and complete fusion. Also, intermediates do occur with high 
frequency in intermediate geographical areas. There is work to be done 
in this regard on the ground in important areas, such as, the Catskill 
Mountains, He Perrot (Montreal) or any other areas where a potential 
close unity of the two appears. 

The past changes in the distribution of the species have been greatly 
exaggerated, especially with regard to any competition between P, rapae 
and P, napi. The habits of the two species are so distinct that no compe- 
tition can reasonably exist. Past records of P, napi feeding on garden or 
cultivated cruciferous plants are undoubtedly correct but these larvae 
came from adults flying out of adjacent woods to lay their eggs on any 
cruciferous plants available. With the destruction of the woods in the 
New England states, the habitat for napi disappeared and then of 
necessity napi disappeared. This was the situation for the low lands. In 
the mountains, napi still exists as it did in the past. 


REFERENCES 

CLARK, A. H. 1951. The butterflies of Virginia. Smithsonian Misc. Col- 
lections, 116(7) : 1-195. 

CLENCH, HARRY K. 1958. The butterflies of Powdermill Nature Reserve. 
Research Report #7. Powdermill Nature Reserve, Carnegie Museum, 
Pittsburgh, Pa. 

EDWARDS, W. H. 1868-1872. Butterflies of North America. Philadelphia. 
The American Entomological Society. 

FISKE, Wm. F. 1901. An annotated catalogue of the butterflies of New 
Hampshire. New Hampshire College, Durham, N. H. 

HOVANITZ, WILLIAM. 1949. Increased variability in populations follow- 
ing natural hybridization. In Genetics, Paleontology and Evolution, 
Princeton. 

— — — — . 1962. The distribution of the species of the genus Pieris in 

North America. /. Res. Lepid. 1 (1): 73-83. 

KLOTS, A. B. 1935. On the life history of Pieris virginiensis Edws. /. N.Y. 
Entom. Soc., 43: 139-142. 

. 1951. A field guide to the butterflies of North America, East 

of the Great Plains. Houghton Mifflin Co., Boston. 

McHENRY, PADDY. 1962. The generic, specific and lower category names 
of the Nearctic butterflies. Part 1. The Genus Pieris. /. Res. Lepid. 
1(1): 63-71. 

SCUDDER, S. H. 1889. The butterflies of the eastern United States and 
Canada. Vols. I-III, Cambridge, Mass., author. 


Journal of Research on the Lepidoptera 1(2) :135-156, 1963 

1140 W. Orange Grove Ave., Arcadia, California, US, A. 

© Copyright 1965 


THE MALE GENITALIA OF SOME COLIAS SPECIES 

BJORN PETERSEN 

University, Lund, Sweden 

In most families of Lepidoptera the genitalia show such 
distinctive characteristics that it becomes, easy to separate even closely 
related species. In the family Pieridae conditions are different. Pieris 
mpi L., P. hfyoniae Ochs., and P. ergane Hbn. are not separable on 
the basis of their male genitalia, nor are F. rapae L. and P. manni Mayer 
(Lorkovic 1928, Drosihn 1933). Similar conditions are present in the 
genus Colias. 

Before going into details a general description of the male genitalia 
of Colias will be given (cf. fig. 1). The distal part of the Vlllth tergum 
is more or less slender, Kusnezov (1915) has called this part the super- 
uncus, Warren (1950) named it the false uncus. At the base of the 
superuncus the tergum is incompletely sclerotized laterally. The saccus 
is rounded, sometimes pointed on its sides. The vinculum is long and 
slender. The tegumen shows a narrow process dorsally, the pseudouncus 
(Kusnezov 1915). The aedeagus has a long ventral arm. 

According to Warren (Lc.) the claspers at their proximal end are 
restricted to a blunt point, attached to the vinculum. The dorsal edge 
is said to be drawn upwards, parallel to the vinculum, and the dorsal 
terminal extremity attached to the tegumen. These statements do not 
correspond to conditions found in Protocolias imperialis (PL I, fig. 1). 
In this species the proximal part of the claspers is rather similar to that 
of other Colias species. The distal part is protracted so that the clasper 
obtains a shape more or less similar to that of many other Pierids. 
It may therefore be concluded that the clasper -head with the terminal 
tooth is the dorsal part of the clasper. 

The short pseudouncus and the marked distal lobe of the clasper 
give to P. impefialis a rather isolated position. In some characteristics it 
comes closer to the genera Catopsilia and Anteos. In other respects, 
however, such as the structure of the clasper head, P. imperialis is 
similar to Colias, thereby showing the relationship between these genera 
(cf.Klots 1929 a and b). 

In the genus Colias the genitalia are rather variable within a species 
or subspecies, as will be shown below. At the same time the differences 
between some species are only slight. Therefore a biometrical approach 
seemed necessary as a complement to the general descriptions of the 


135 


136 


PETERSEN 


J. Res. Lepid. 


genitalia. The modes of measuring and the results are given first as 
important for the full understanding of the descriptions. 

The breadth of the superuncus. The edges of the superuncus in 
dorsal view may a) converge distally b) run parallely or c) diverge 
slightly from a narrow part near the base ( cf . fig. 2 ) . The breadth was 
on diverging superunci measured at the narrowest basal point, on 
others at the corresponding place. 



Fig. 1. The male genitalia of a Colias. a superuncus b pseudouncus 
c clasperhead d tooth of clasperhead e uncus 

The size of a certain part of an individual may be influenced by its 
general size. The breadth of the superuncus has therefore been correlated 
with a measure of size: the wing length. 

All species with genitalia similar to those of the three species 
previously investigated: C. hecla, nastes and palaeno are included in 
fig. 3. Here the breadth of the superuncus is plotted against the wing 
length in a logarithmic scale. The populations investigated are parted in 
two groups, the first with a slender superuncus, the second with a broad 
one. The regression line of the populations with a slender superuncus 
is: 

log y 1 = 0.75 log X + 0.68; (r = 0.72; 0.02 >£> 0.01). 

The species along this regression line are both orange with a hecla- 
pattern, yellow with a ^^c/^-pattern, and yellow with a nastes-patitm. 
The species in fig. 3 with a broad superuncus are either orange with a 
hecla-^RttQm or yellow with a nastes-pRttQm. These two types of species 
can be grouped along two regression lines. Within both groups the 
correlation is statistically significant. 


t(2):i}5-i56, 196 } 


COLIAS GENITALIA 


137 


Orange species: 

log y=: 0.80 log X + 0.77; (£ = 0.71; 0.05 > P > 0.02). 
Yellow species: 

log y 1.17 log X + 0.30; (r = 0.81; 0.05 > P > 0.02). 



Fig. 2. Differently shaped superunci with semicircular, less sclerodzed areas 
near the base., a convergent b parallel c divergent 


The means of the two groups are, however, not significantly separated. 
An analysis of covariance gives 0.1 > ^ > 0.05. 

The data on which fig. 3 have been based are tabulated in Tables 1 
and 2. Among the species shown in these tables C. nastes is of special 
interest. In Scandinavia and in North America but also in Altai and 
the Sayan Mountains the superuncus is slender. C. nastes of Siberia 
and the Amur Province is intermediate between nastes of the Altai and 
the Sayan Mountains and the closely related, allopatric C. montium from 
S. Kansu. This species forms a transition to C. cocandica from Ferghana, 
the Issykkul, and from the Tianshan. The superuncus-breadth of all 
Asiatic individuals of nastes is plotted against wing length in fig. 4, 
The populations are grouped along two regression lines, though the 
correlation in none of the groups is statistically significant ( cf . Table 1 ) . 
An analysis of covariance, however, shows that the means of the two 
samples (shown in the figure by squares) are significantly different 
(0.01 > P> 0.001). 

In the yellow series of forms there is thus no sharp limit between 
species with slender and broad superunci. Even within a single popula- 
tion, that from the Amur Province — the variation is so great that it 
includes superunci typical f(Jr nastes nastes and others typical for 
cocandica (cf. fig. 4). In the orange series of forms conditions are 
different as will be treated later. In fig. 5 the superuncus breadth of 
arctic C. nastes and of its sibling in the Alps, C. phicomone, is plotted 
against wing length. The differentiation has proceeded so far that 
hardly any overlap is present. 

The wing length and breadth of the superuncus in some other 
Colias species, less similar to those treated above, are shown in Table 3. 
In the three pairs at the top of the table the genitalia are so similar that 


138 PETERSEN /• Lepid. 



KOU J L I I ! I ( 1 I 

1.30 1.32 1.34 l36 1.38 1.40 1.42 7.44 7.46 7.48 


Fig. 3. Superuncus breadth of some Colias populations plotted against wing 
length. Data from Tables 1 and 2. 

O Population from N. America or northern Eurasia 
□ Population from Central Asia or Central Europe 
7\ C. palaeno and interior 

rod 3hoNe=hecla patterns open mark=orange color 

rod helow—nastes patterns filled mark=yellow color 


no constant differences could be found. Warren (1950) states some 
differences between C. australis and hyale, a statement which was not 
confirmed by the specimens investigated. 

Two of the species of Table 3, C. cunninghami and P. imperialis, 
have an extremely broad superuncus. C. vautieri, which otherwise is 


1 ( 2 ): 135 ^ 156 , 

1.90 ^ 


.?«3 COLIAS GENITALIA 

log superuncus breath 


1.85 


O 




139 


1.80 




1.60 



o + 


log wing length 

U. I -J_ — __ 

1.32 1.34 1.56 1.38 1.40 1.42 1.44 

Fig. 4. Superanciis breadth of some Asiatic populations of C. nastes plotted 
against wing length. The lower square represents the mean of the dots, the 
upper square the mean of the remaining marks. 

• Altai and Sayan Mts. A Transbaikal 

O Siberia, Taiga zone + Amur Province 

rather aberrant, C. erate, and myrmidone fit well into the group of 
species with a broad superuncus, listed in Table 2. The remaining 
four species have a more slender superuncus though C. hyale and 
australis, like C. montium occupy a somewhat intermediate position. 
The form of the superuncus in lateral view-. During copulation 


PETERSEN 


/. Res. Lepid, 


140 

Log superurunijS breadth 

100^ 


95 - 


o 

o 


o 

o 


90 - 


85 - 

80 - 


O 



O 


O 


O 

o oo 


o 


o 


o 


o 


o 


o 


o 


75^ 



65 - 


60- 


55 - 

126 1.50 tsz 134- 1.56 


log wing Length 
138 UO 142 


Fig. 5 Superuncus breadth of C. phicomone (open circles) and C. nastes 
(filled circles = Scandinavia, + North America) plotted against wing length. 


(2): i }5~ i 56, 1963 


COLIAS GENITALIA 


.141 


the superuncus together with the uncus is pressed against the body of 
the female, just as the uncus in the genus Pieris according to Lorkovic 
(1947). To make possible the bending down of the superuncus during 
copulation a semicircular area laterally on the Vlllth tergit is less 
sclerotized than the rest ( cf . fig. 2). This bending can be stated without 
studying any copulating pairs. In some specimens the superuncus is 
broken (cf. fig. 6 ), in some in addition kept between the two claspers. 



Fig. 6. Colias intenor with superuncus broken during copulation. 


The breadth of the superuncus of such specimens sometimes cannot 
be measured without maceration. A broken superuncus was found in 
all species where more than a few specimens were investigated, as 
shown in Table 4. 

Only in very few species the superuncus is a straight process 
protruding from the Vlllth tergit. Also when unbroken the superuncus 
usually forms a bow downwards. This bow is present, even in its most 
pronounced form, in animals which have not copulated, as, for instance, 
in all the thirty C. hecia specimens investigated which in 1952 were 
caught at the beginning of the flying time on the northern side of 
the Lake Tome Trask. 

To obtain a quantitative estimation of the variation of the form of 
the superuncus in lateral view, the angle of the distal part of the 
superuncus to the dorsal edge of the Vlllth tergum was measured. 
The result of this investigation is shown in Table 4. In general a 
broad superuncus is only slightly bent downwards while a narrow is 
bent more. The exceptions are rather few: C. interior, montium, and 
phicomone. 


142 


PETERSEN 


/. Res. Leptd. 


Species 

n 

log X 

M, 

log y 

r 

P 

b 

A. Orange species 







C, hecla, Scandinavia 

51 

1.331 

1.70 

-0.18 

0.3-0. 2 

-0.74 

, Siberia, Dudinska 

4 

1.355 

1.67 




, Baffin, Isl. 

2 

1.347 

1.79 




, Greenland 

3 

1.362 

1.74 




C, hyperborea, N. Siberia 

4 


1,69 




C. eurytheme, N. America 

6 

1.456 

1.76 




, slightly orange 
Texas, Houston 

6 

1.313 

1.61 




B. Yellow species 







a. hecla pattern 







C. palaeno, Europe 

41 

1.411 

1.71 

0.18 

0.3-0. 2 

0.50 

C. palaeno, Siberia 

9 

1.377 

1.71 

0.57 

0. 2-0.1 

1.73 

C. interior 

2 

1.311 

1.67 




b. nastes pattern 







C, nastes, Scandinavia 

42 

1.344 

1.68 

-0.09 

0.6-0. 5 

-0.24 

, Siberia, Amur area 

29 

1.391 

1.75 

0.32 

0.1-0.05 

0.87 

, Altai, Sayan Mts. 

12 

1.379 

1.67 

0.35 

0.3-0. 2 

0.86 

, N, America 

2 

1.311 

1.68 




C. philodice, N. America 

11 

1.419 

1.74 




C. montium, S, Kansu 

9 

1.370 

1.77 





Table 1. Wing length (s<), in mm, and breadth of superuncus (y), in an arbitrary scale. 


and their correlation in some Colias species with a slender superuncus. Logarithmic 
scale. 

C. interior is in this character well separated from its allopatric 
sibling, C. palaeno, in having a rather straight but slender superuncus. 

C. montium has a rather slender superuncus which is straight as is 
that of cocandica. In this respect another of the southern species, 
C, phicomone, is intermediate between nastes and cocandica. The 
superunus of phicomone is, however, rounded and not more or less 
pointed as it is in all other species with a slender superuncus except 
C. interior, aurorina, and sagartia. 

By means of the two characteristics, i.e. the breadth and the 
form of the superuncus, it is possible to separate the two species 
C. viluiensis Men. and hyperborea Gr. Gr. which live sympatrically in 
N.E. Siberia. Of viluiensis 7 specimens from the Verchojansk area, 
Lutsha near Yakutsk, Vilutsk and Vilui have been investigated. The 
four specimens of hyperborea were from Sib. poL, the Lena Valley 


I ( 2 ) ; I )5-i56, 7963 


COLIAS GENITALIA 


143 


Species 

n 

”log X 

M, 

log y 

Species 

n 

'^log X 

M 

log y 

Orange species, 

hecla-pattern 




Yellow species, 

nastes-pattern 




C, chrysotheme 

5 

1.350 

1.89 





C. croceus, Europe 

10 

1.403 

1.90 

C. alpherahyi 

1 

1.436 

2.02 

, Asia 

11 

1.429 

1.89 

C. christophi 

1 

1.369 

1.86 

C. eogene 

8 

1.368 

1.87 





C. heos 

12 

1.474 

1.99 

C. cocandica 

2 

1.326 

1.85 

C. remanovi 

2 

1.446 

1.91 

C. phicomone 

24 

1.374 

1.88 

C. standingeri 

6 

1.395 

1.83 

C. sieversi 

1 

1.405 

1.93 

C. thisoa 

2 

1.417 

1.90 

C. sifanica 

1 

1.326 

1.90 

C. wiscotti 

9 

1.431 

1.93 





C. viluensis 

7 

— 

1.90 






Table 2. Wing length (x), in mm, and breadth of superuncus (y), in an arbitrary 
scale, ih some Colias species with a broad superuncus. Logarithmic scale. 

and Sredne Kolymsk.. The latter specimens all have a slender and 
strongly bent superuncus of the hecla-type, while all specimens of 
viluiensis have a straighter and broader superuncus. There is no overlap 
in any of the two characters; in the "angle-character” the gap is very 
wide. 

In Colias nastes from Asia there Is no similar correlation between 
the breadth and the shape of the superuncus. Among the 41 specimens 
investigated the coefficient of correlation is -j- 0.093 which is far below 
significance ( 0.8 > P > 0.7 ) . 

Conditions are thus quite different in northern Asia as regards the 
nastes- and hecla-stn^s of forms. In the nastes-SQnts dines including 
characters of the genitalia reach the Amur area, Transbaikal, the Sayan 
Mts, and the Altai, and forms intermediate between arctic and Central 
Asiatic ones are present in North Western China and in the mountains 
of Central Europe. In the orange series of forms an overlap of a northern 
and a southern species is present between 65° -68° n. latitude and no 
intermediates have yet been found between these species. 

Number of teeth near apex of aedeagus. In many groups of insects 
the number, shape, and position of the aedeagal teeth serve as good 
evidence to distinguish species. In the Colias species all the teeth are 
small, of a rather similar shape, situated near the apex. Variation in 
number is strong even within subspecies when compared with the 
differences occurring between species. Thus this character is without 
signifiance for the determination of individuals. A closer investigation 
of a material belonging to the group of species with a slender super- 


144 


PETERSEN 


/. Res. Lcpid. 


C. aurorina 

n 

M, 

log X 

M, 

log y 

6 

1,449 

1.80 

C, sagartla 

5 

1.437 

1.76 

C. australis 

8 

1.384 

1.74 

C. hyale 

27 

1.373 

1.79 

C . erate 

13 

1.387 

1.94 

C, myrmidone 

7 

1.425 

1.95 

C. cunninghami 

2 

1.367 

2,01 

C. vautieri 

2 

1.327 

1.03 

P. imperialis 

1 

1.356 

2.05 


Table 3. Wing length (x), in mm, and breadth of superuncus (y), 
in an arbitrary scale, in some Colias species. 

uncus, however, revealed some slight specific differences in the average 
number of teeth (cf. Table 5). The results of this investigation are; 

C. nastes has on an average fewer teeth than C. hecla from the 
same locality (Scandinavian material t = 2.45; 0.02 > P > 0.01, 
American material not differing significantly. There is a certain tendency 
of parallelism, both species having a lower number of teeth in Scandi- 
navia than in North America. This tendency is not significant in any 
of the species, however. C. palaeno and interior both have a low number 
of teeth, while the number is fairly high in C. meadi, philodice, and 
eurytheme. 

SPECIAL PART 

The genus Colias may be parted in two genera: Protocolias (type 
imperialis Btlr.) and Colias, the latter in turn in two subgenera 
Mesocolias (type vautieri Guer.) and Colias. The descriptions of the 
genera, the subgenera and their various species may be given most 
easily in the form of a key. 

1. Small or medium-sized, orange butterflies with broad (0.40-0.45 
mm) or very broad (0.5 mm), straight superunci. Superficial scales 
— if present— -yellow and black, broad and flattened in the distal 
part. Apex of aedeogus without teeth. 

Genus Protocolias and subgenus Mesocolias 2. 
r. Small to big, yellow, greenish or orange species with broad to 
slender (0.32 mm) superunci. Superficial scales in the black mar- 
gin— if present— yellow, pointed, hairlike or broader. A number 
of teeth at apex of aedeagus 

Subgenus Colias 3. 


i(2):i}5-i56, 196 } 


COLIAS GENITALIA 


145 


2. Superuncus very broad and triangular. Pseudouncus short. Clasper- 
head with elongated tooth. Middle part of the clasper with a 
marked prong. Ventral arm of aedeagus broader in the distal end. 
Black and yellow suoerficial scales present. 

Pfotocolias imperialis Btlr. (PI. 1:1) 

2’. Superuncus broad. Tooth of clasperhead slightly bent upwards. 
Ventral lobe (v.l.) of inner side of the clasper (Klots 1929 a) 
more marked than in any other species of the genus. Distal part of 
ventral arm of aedeagus showing charaaeristic shape. Superficial 
scales absent. 

Colias (Mesocolias) vautieri Guh. (PI. 1:2,3) 

2”. Superuncus very broad, genitalia small, claspers short ( 1.0-1. 1 mm) 
but broad (0.6 mm). Distal part of ventral arm of aedeagus not 
broader than the proximal one. Superficial scales absent. 

Colias {Mesocolias) cunninghami 'Qth. (pi. 1:4). 

3 . Claspers caudally with a marked prong directed medially. Super- 
uncus straight and fairly slender. 

C. hyale L. (pi. 1:5) and australis Ver. 

3’. Claspers without any marked prong ........4. 

4. Claspers pointed in the middle part of the caudal edge 

(most easily seen from behind) ........ 5 . 

4’. Claspers not pointed in the middle part of the caudal edge 6. 

5. Superuncus straight, on an average 0.40-0.45 mm broad 

C. erate Esp., C. myrmidone Esp. (PI. 1:6), 

5’. Superuncus usually markedly bent downwards, 0.30-0.35 mm broad. 

C, sagartia Led. ( PI. 1 : 7 ) C. aurorina H. Sch. (PI. 1:8). 

6. Superuncus on an average less than 0.32 mm broad (cf. text fig. 3) 

usually strongly bent downwards (cf. Table 4) ......... 7 . 

6’. Superuncus on an average broader than 0.35 mm, usually 

straight 13. 

7. Superuncus only slightly bent downwards .......... 8 , 

7’. Superuncus strongly bent downwards 9. 

8. Superuncus in caudal view blunt-ended as in species with a broad 
superuncus. Inner side of -clasper of hecla-type (cf. PI. 11:34) 

C. montium Oberth. (PI. 1:9, 10). 

8’. Superuncus in caudal view pointed. Inner side of clasper with well 
developed ventral lobe and ridged from its dorsal edge dorso- 
caudally towards the caudal part of the clasper (Fig. 14, 15) 
C. interior Scudd. (PI. I: 11-13, II: 14, 15), 

9 . Ridge from dorsal edge of ventral lobe towards caudal part of the 
clasper present as in C. interior. Ventral lobe less well developed 
than in C. interior (cf. Fig. 16). Caudal edge of clasper often 
strongly bent inwards ( most easily seen in caudal view ) . 

C. palaeno L. (PI. 11:16-25). 

C. Christina (PI. 11:26-28). 
C. palaeno: is most variable in all charaaers investigated except 
the breadth of superuncus (cf. the figures). One specimen from 
Abisko (42 specimens studied) completely lacks the ridge on the 
inner side of the clasper. Hence it cannot with certainty be separ- 
ated from C. nastes or hecla. The limited material of C. Christina 
appeared rather close to some specimens of palaeno, though it 
might be possible to separate the two species after examining a 
greater material. 

9’. The caudal edge of the valva less strongly bent inwards ventrally. 
Inner side of clasper reminding of hecla-type (cf. fig. 34) ........10. 

10. Clasoer (in lateral view) broadest rather ventrally 11. 

10’. Clasper broadest in the middle part 12. 

11. Aedeagus on an average with a higher number of teeth at the distal 

end ........................................C hecla Lef. (PI. 11:29-39). 


146 


PETERSEN 


/. Res. Lcpid. 


Frequency of 
superuncus 
broken 


7/50 






1 14/56 1 

1 14/56 1 

14/56 

1/12 

4/25 

Ipdl 1 


1 1 

2/9 1 



2/12 


<D 

DO 

C 

< 



82 

s 

s 

cr> 

ID 

s 

s 


<T> 

CD 

s 

O 


to 

r- 

.It 

CO 

El 



CM 


=t- 

o 





- 
















95-104 1 


CM 


- 

c. 


- 





*— J 

if 


- 








- 


CO 

CO 

- 

J- 

- 







- 






3 





- 

- 

*— 1 

- 

•— J 

- 


CD 

- 

f—{ 

CO 






d- 

S 







- 

- 

- 

- 




- 

- 






ID 


- 







- 

it 


*— 1 

CM 

- 




- 


- 

3 

m 

a- 



- 


- 



rH 


fH 

o 

c 

- 







CM 




#— J 






- 


- 





CO 


- 



25-341 











tD 








- 

- 

CN 

1 








•H 











CM 

•— 1 

it 

•H 





















Species, 

Population 

Popu of Table 1. 

C. hecla, Scand. Mts. 

" , N. America 

C. hyperborea 

C. meadi 

j C. eurytheme | 

C. palaeno, Scand. Mts. 

j ” , Europe | 

1 " , Asia 

" , N, America 

C. interior [christina] 

m 

■p 

'O 

c 

o 

CD 

(0 

«D 

CO 

m 

c 

m 

CO 

< 

" , N. America 

1 C. philodice 

1 C. beryl la 

Pop. of Table 2. 

C. chrysotheme 

i 

C. croceus, Europe 

0} 

c 

0) 

DO 

0 

o 


IT. Aedeagus on an average with a lower number of teeth. Very 
similar to C. hecla .....C, nastes Bdv. (PI. III:35"38). 

12. Bigger, superuncus broader ......C. eurytheme Bdv. (PL 111:39, 40). 

C. philodice Gdt. (PI. III:4l). 

12’. Smaller, superuncus slenderer. ....C. meadi Edw. (PL 111:42, 43). 

13. Superuncus bent downwards C. phicomone Esp. (PL 111:44). 

14. Superuncus diverging, very broad near the distal end. Clasper (in 

lateral view) broadest ventrally C. croceus Fourcr. (PL 111:45). 

14’. Superuncus often with parallel or converging edges or clasper 
broadest in the middle part 15. 

15. Clasperhead more strongly marked, tooth strongly developed. Ven- 
tral arm of aedeagus broad in the distal part. 

C. wiscotti Stgr. (PL 111:46, 47) 
C. marcopolo Gr. Grsh. probably also belongs here. No material 
has been available for investigation. Professor Sheljuzhko informs 


I (2); 1 ) 5 - 156 , 196) 


COLIAS GENITALIA 


147 


2/10 I 


CO 













1 

4/30 

2/15 





0 

0 

0 

fO 

0 

CD 

CM 



0 

fO 

0 

CO 

CD 

to 

0 

0 



s 

CO 

CO 

o 

. 1 ^ 

o 

CO 

o 

CM 

o 






















































zS- 






















- 




e. 

CM 






















f—h 








- 




- 





c 






- 

CM 


- 













CO 







CM 






- 




eH 

iH 








- 

- 

- 







- 




CM 

CO 






- 




CO 


- 

- 




OJ 




CO 

CM 





rH 




t—i 


CM 


- 








- 













- 




- 

j C. heos 1 

C. romanovi 

C. staudingeri 

C. thisoa 

C. wiscotti 

1 C, vlluensis | 

j C, alpherakyi j 

C. christophi 

j C. cocandica 

i 

B 

0 

0 

U 

j C, sieversi | 

C. sifanica 

Pop. of Table 3. 

C. aurorina 

C, sagartia 

j C« australis | 

C. hyale 

C. erate 

j C* rayrmidone j 

C. cunninghami 

! C. vautieri | 

j P. imperialis j 


that Avinov and Kusnezov both place marcopolo together with 
wiscotti, the latter as a subspecies. According to Sheljuzhko both 
forms fly together in the Pamir area and are probably different 
species in spite of a great similarity in the genitalia. 

15 ’. Clasper head less strongly marked 16. 

16. A number of species remain which due to lack of sufficient 
material and/or great similarity could not be separated from each 
other : 

Yellow species with nastes-p2XtQtn\ C. dpherakyi Stgr. (PI. 
111:48), C. chfistophi Gr.-Grsh. (PL IV:49), C, cocandica Erscii. 
(PI. IV: 50) (Specimen figured with triangular saccus; in a second 
specimen investigated the saccus is rounded as in all other species), 
C, sieversi Gr.-Grsh. (PI. IV:51), C. sifanica Gr. Grsh. (Pi. 
IV: 52). These Central Asiatic species together with C. nastes, 
phicomone, montium and others form one or — most probably — 
two polytypic species. 


148 


PETERSEN 


/. Res. Lepid. 



n 

2 

3 

4 

5 

6 

7 

0 

9 

10 

11 

12 

M 

C. hecla, N. Am. 

8 





5 

1 

2 





6.6 

*' , Greenl. 

3 



1 

1 

1 






5.0 

" , Scand. 

30 



2 

7 

9 

8 

4 





6.2 

C. meadi 

10 





4 

1 

3 

1 



1 

7.6 

C. eurytheme 

12 




3 

3 

5 


1 




6.4 

C. palaeno, N. Am. 

10 


1 

1 

5 

2 


1 





5.2 

'* , Asia 

8 


1 


1 

6 







5.5 

" , Estonia 

8 


1 


3 


2 

1 

1 




6.1 

*' , Scand. 

21 

1 

2 

7 

2 

3 

1 

3 

1 

1 



5.4 

C. interior 

11 



3 

4 

2 

2 






5.3 

C, Christina 

1 





1 







6 

C, nastes, N. Am. 

6 




1 

4 


1 





6.2 

*' , Asia 

5 




1 

3 

1 






6.0 

” , Scand. 

23 


1 

4 

8 

5 

5 






5.4 

C. philodice 

5 




1 

2 

1 




1 


7.0 


Table 5, Number of teeth near the apex of the aedeagus of Colias species with a slender 

superuncus. Orange species with hecla-pSLttern: C, chrysotheme Esp. (PI. 

IV:53), C, eogene Fldr, (PI. IV; 54, 55), C. staudingeri Alph. 
(PI. IV: 56 ), C. thisoa Men. (PI. IV:57), C. viluiensis Men., 
and C. heos (PI. IV:58, 59). As in the previous group it is un- 
certain whether all these forms deserve each a specific status. 

Orange species with «^j/^j-pattern : C. romanovi Gr. Grsh. 
(PL IV:60). 

This enumeration of Colias species is not complete. 


DISCUSSION 

The key given above does not show the phylogenetic relationships. 
These are, due to extensive introgressive hybridization, very difficult 
to find out. Suppose species A is closest related to the allopatric species 
B which has given rise to species C sympatric to A. Due to the 
introgressions A C or C A it is then possible that A and C 

are more similar in several characters than A-B or B-C. However, some 
characteristics, as for instance those on which the isolation between A 
and C depend, are likely to be of maximum difference in the case A-C. 

Among the Co/m-species investigated there exists a number of 
pairs of species within which the male genitalia are very similar: 
C. eurytheme-philodice; C. hecla-nastes; C. aufofina-sagartia; C. myrmi- 
done-erate; C. hyale-australis and probably also C. pdaeno-christina and 
C. wiscotti-mar copolo. Introgression has among these been established 
between C. eurytheme-philodice (Hovanitz 1949 a, b) and between 
C. hecla and nastes. Hybridization has been suggested between C. sugar- 
tia and aurofina (Lederer 1941). It is possible that pairs of a similar 
kind are present in Central Asia among the species figured from number 


(2); i }5- i 56, 1963 


COLIAS GENITALIA 


149 


48 to number 58. Lederer (1941) mentions, among other suspected 
hybrids, specimens which have been supposed to be hybrids between 
C. eogene and cocandica. A number of species with different genitalia 
have been seen in copula ( Lederer l.c. ) : hyde x myrmidone, hyde x 
croceus, hyde x erate, croceus x erate, and hyde x phicomone. The last 
cross gave rise to some larvae which died before the pupation. All 
these species have markedly different genitalia and are probably not 
very closely related. It therefore seems uncertain whether hybridization 
in these cases can give rise to introgression. If introgression occurs it 
must be possible to observe this fact also on the . genitalia. 

In all the pairs of sibling species first mentioned {eurytheme- 
philodice etc.), except hyde-austrdis and pdaeno-chfistina, colors, and 
in most cases also patterns, are different. The first species is usually 
orange with a heda-p 2 XXQm, the second yellow with a nastes-^ 2 XX^iVi. 
Only C. sagarda has in addition a blue pigment which is present in 
some individuals of its orange sibling aurorina. 

The common occurrence of differences in color and pattern between 
sibling species suggests that these colors are integrating part of the 
isolating mechanisms within the pairs. Sexual isolation of this kind has 
in the pair Pieris napPbryoniae been established by Petersen, Tornblom 
and Bodin (1952). Males of both species are attracted by the white 
color of the P. napi female. The yellow female of bryoniae attracts the 
males solely by means of movements and odors. No releasing effect of 
any of the types of pattern was obtained neither in these pierids nor 
in similar experiments with the Silverwashed Fritillary {A. paphia L.) 
(Magnus 1954). Different color but probably not different pattern may 
therefore play a role for the sexual isolation between Colias species. 

The geographical distribution of some subdivisions of the genus 
Colias may also be discussed. Protocolias and Mesocolias are entirely 
South American. The species of the subgenus Colias with a broad 
superuncus are all Palearctic, one group having penetrated even into 
the Ethiopian region. Colias with a slender superuncus are mainly 
distributed in North America and the northern Palearctic, only a few 
living further south in the latter region. 

The distribution of the groups is to a great extent certainly the .. 
result of an evolution within different areas. It does not seem established 
that palearctic Colias have evolved from South American forms. They 
may as well have developed from primitive Colias in some other part 
of the world where they are now extinct. 

The evolution of the superuncus. Among the two characteristics 
of the Colias genitalia, the superuncus and the pseudouncus, the latter 
is present in the genus Anteos as well as in Catopsilia and Colias (Klots 
1929 a, b, Drosihn 1933). The superuncus, on the other hand, is very 
small and triangular in the Anteos species investigated {menippe, 
clofinde, Plate IV, figs. 61, 62) covering only the pseudouncus and the 
basal part of the uncus. A similar, though bigger superuncus is found 
in Protocolias imperialis and the genus Phoebis (Drosihn 1933 ). In 


150 


PETERSEN 


/. Res. Lepid. 


the latter genus no pseudouncus is present. The Catopsilia and Aphrissa 
species have broad, straight and diverging superunci (cf. Plate IV, 
figs. 63, 64 and Drosihn 1933), rather similar to those of some Colias 
species. The triangular shape of the superuncus is probably primitive, as 
this shape is present in Anteos where the superuncus is comparatively 
small. 

The superuncus has in the genus Colias (and probably also in 
Catopsilia and Aphrissa) taken over the function of the uncus of many 
other Lepidoptera (cf. Lorkovic 1947) to assist medially and dorsally 
in holding the female body during the copulation. The superuncus 
and the pseudouncus probably developed to support the uncus dorsally. 
As the superuncus became larger, it was placed directly against the 
female body, and thus the uncus instead changed to support the super- 
uncus. 

It has been suggested, for instance by Verity (1947), that Colias 
have developed from the Old World Catopsilias. The presence of the 
very primitive Protocolias with a triangular superuncus, a short pseudo- 
uncus, and the middle part of the clasper with a prong, makes this 
assumption rather unlikely. The triangular superuncus may be con- 
sidered as a very pronouncedly convergent one ( cf . fig. 2 ) . The more 
or less parallel or divergent superunci of Colias, Catopsilia and Aphrissa 
may have evolved by means of parallel evolution towards more uncus- 
like conditions as discussed below in the case of Colias. 

The Catopsilias and most Colias species have broad, straight, and 
unpointed superunci, characteristics which therefore may be considered 
primitive compared with those of C. hecla, nastes and palaeno. A 
similiar result is arrived at, if the problem is approached from another 
direction. As already mentioned the superuncus has in the genus Colias 
taken over the function of the uncus. The latter has had its function 
during so much longer a time that it may give a certain indication of 
what shape is most apt to give an optimal function. The uncus is 
slender, pointed, and rather straight. In the two first of these character- 
istics the uncus corresponds to the superuncus of the hecla-nastes- 
palaeno-x^^t. Only it is even more pronounced then. The shape of the 
uncus on the other hand, is straighter in lateral view. The basal part of 
the superuncus is, however, situated more dorsally than the same part 
of the uncus ( cf . textfig. 13). Only if the superuncus is bent down- 
wards, it can be placed against the female body in the same place as 
the uncus of other Pierids. 

The evolution of the superuncus thus seems to have converged 
with that of the uncus. There is, however, still a marked difference in 
shape between these two organs and for different reasons it is not likely 
that the convergence will become complete ever. As already mentioned 
the situation of the two organs is different. The pressure of the super- 
uncus is supported by that of the uncus and probably also by that of 
the pseudouncus, and finally the claspers are in the genus Colias not 
built as in other genera. 


( 2 ): i ) 5 - i 56 , 1963 


COLIAS GENITALIA 


151 


SUMMARY 

The male genitalia of a number of Colias species have been de- 
scribed. The genus is divided into one new genus and two subgenera 
of the genus Colias: The South American genus Protocolias (type 
imperialis Btlr.) and subgenus Mesocolias (type vautieri Guer.) and 
the mainly Holarctic subgenus Colias. The latter can be divided into a 
palearctic group with a broad straight superuncus and a nearctic and 
northern palearctic group with a slender superujicus which is bent 
downwards. Transitions between these two groups exist within the 
supraspedes C. nastes. 

The evolution of the superuncus from a small beginning, as at 
present in the genus Anteos, via the broad superund of the Catopsilias 
and some Colias into a slender superuncus has been discussed. 

Several pairs of Colias species with identical or very similar geni- 
talia exist. It is suggested that the similarity is combined with intro- 
gression. Several of the pairs include species of different colors. These 
colors may serve to isolate the species sexually from each other. 


REFERENCES 

DROSHIN, J- 1933. Uber Art-und Rassenunterschiede der mannlichen 
Kopulationsapparate von Pieriden, 1-135.. Stuttgart. 

HOVANITZ, W. 1949a. Increased variability in populations following 
natural hybridization. In Jepseo, et al. Genetics, Paleontology, and 
Evolution. 339-355. Princeton. 

HOVANITZ, W. 1949b. Interspecific matings between Colias eurytheme 
and Colias philodice in wild populations. Evolution. 3: 170-173. 
KLOTS, A. B. 1929a. The genus Anteos Hubner ( Lepidoptera, Pieridae). 
Bull. Brooklyn Ent. Soc. 24: 134-214. 

^ 1929b. The generic status of Catopsilia Hubner and 
Phoebis Hubner, with a discussion of the relationships of the species and 
the homologies of the male genitalia (Lepidoptera, Pieridae). Bull. 
Brooklyn Ent. Soc. 24: 203-214. 

KUSNEZOV, N. J. 1915. Insectes Lepidopteres I in Faune de la Russie 
1-337. Petrograd. 

LEDERER, G. 1941. Die Naturgeschichte der Tagfalter. Stuttgart. 
LORKOVIC, Z. 1928. Analyse des Speziesbegriffes und der Variabilitat der 
Spezies auf Grund von Uotersuchungen einiger Lepidopteren. Glasn. 
Soc. Scient. Nat. Croat. 38: 1-64, 

LORKOVIC, Z, 1947. Modes artifidels d’accouplement des papillons. 
Glasn. Biol. Sekc. Ser. II/B 1 :86-98. 

MAGNUS, D. 1954. Experimentelle Untersuchungen am Kaisermantel zur 
Analyse optischer Auslosungsreize. Deutscher Entomologentag in Ham- 
burg 30. Juli bis 3. August 1953: 58-75. 

PETERSEN, B., O. TORNBLOM, and N. -O. BODIN. 1952. Verhaltens- 
studien am Rapsweissling und Bergweissling (Pieris napi L. und Pieris 
bryoniae Ochs.) Behaviour. 4: 67-84. 

WARREN, B. C. S, 1950. Speciation in the genus Colias: with special 
reference to C. hyale and C. australis. Lambillionea. 50: 90-98. 


152 


PETERSEN 


/. Res. Lepid. 


PLATES 

a. lateral view 

b. superuncus, dorsal view 

c. caudal view 

d. aedeagus, ventral or dorsal view 

e. ” , distal end in lateral view 

f. clasper from inner side 

g. saccus in ventral view 
u. uncus 

v.l. ventral lobe of the inner side of the clasper (Klots 1929 a) 


PLATE I. 

Fig. 1. P. imperialis (wrongly labelled 





Honolulu) 

Fig. 

2. 


C. vautieri, Ensenada 

Fig. 

3. 


C. vautieri, Ensenada 

Fig. 

4. 


C. ctinninghami, Junin, Peru 

Fig. 

5. 


C. hyale, Oland, Sweden 

Fig. 

6. 


C. myrmidone , Germany 

Fig. 

7. 


C. sagartia, N.E. Persia 

Fig. 

8. 


C. aiirorina, Armenia 

Fig. 

9. 


C. montium, Tatsienlou, Tibet 

F.g. 

10. 


C. moiitium, S. Kansu, China 

Fig. 

11. 


C. interior, Montreal, Canada 

Fig. 

12, 

13. 

C. interior, Alaska Highway, mil 
126, Beatton R. area, B.C. 

PLATE 

II. 


Fig. 

14. 


C. interior, Alaska Highway, mil 
90, Beatton R. area, B.C. 

Fig. 

15. 


C. interior, Alaska Highway, mil 
90, Beatton R. area, B.C. 

Fig. 

16. 


C. palaeno, Alaska Highway, mil 
450, Toal River, B.C. 

Fig. 

17. 


C. palaeno, Bjurfors, Sweden 

Fig. 

18. 


C. palaeno, Sweden 

Fig. 

19, 

20. 

C. palaeno, Smaland, Sweden 

Fig. 

21. 


C. palaeno, Abisko, Sweden 

Fig. 

22, 

23. 

C. palaeno, 10 miles South 

Burwash Landing, Y.T. 

Fig. 

24, 

25. 

C. palaeno, Sweden 

Fig. 

26. 


C. Christina, America borealis 

Fig. 

27. 


C. Christina, Rocky Mts. 

Fig. 

28. 


C. Christina, Am. bor. 

Fig. 

29. 


C. hecla. Nr. Haines Junction, 
Y.T., up Summit Cr. 

6000’-7000’ el. 

Fig. 

30. 


C. hecla. Bog. nr. Johnson’s 
Crossing, Y. T. 

Fig. 

31. 


C. hecla, Dudinska, Siberia 

Fig. 

32. 


C. hecla, Sweden 

Fig. 

33. 


C. hecla, Kvikkjokk, Sweden 


Fig. 34. C. hecla, 20 miles South 
Burwash Landing, Y. T. 

PLATE III. 


Fig. 

35. 


C. nastes. Nr Haines Junction, 

Y. T., up Summit Creek, 

6000’-7000’ el. 

Fig. 

36. 


C. nastes, Mt. Atabaska, Jasper 
N.P., Alberta, 7000’-8000’ el. 

Fig. 

37, 

38. 

C. nastes, Kvikkjokk, Swedish 
Lapland 

Fig. 

39. 


C. eurythemc, Texas 

Fig. 

40. 


C. eury theme , Minnesota, U.S.A. 

Fig. 

41. 


C. philodice, Amer. bor. 

Fig. 

42, 

43. 

C. meadi. Bow Pass, Jasper N. P., 
Alberta 

Fig. 

44. 


C. phicomone, Alps 

Fig. 

45. 


C. croceus, Tirol, Austria 

Fig. 

46. 


C. wiscotti, Turkestan 

Fig. 

47. 


C. wiscotti separata, Turkestan 

Fig. 

48. 


C. alpherakyi , Turkestan 

PLATE 

IV. 


Fig. 

49. 


C. christophi, Turkestan 

Fig. 

50. 


C. cocatidica, Turkestan 

Fig. 

51. 


C. sieversi, Turkestan 

Fig. 

52. 


C. sifanica 

Fig. 

53. 


C. chrysothemc 

Fig. 

54. 


C. eogene, Kisil Fast area 

Fig. 

55. 


C. eogene 

Fig. 

56. 


C. staudingeri, Tian Shan, 
Fu-Shu-Shan 

Fig. 

57. 


C. thisoa 

Fig. 

58. 


C. heos, N. Mongolia, long. 100°- 
110°, lat. 45°-50° 

Fig. 

59. 


C. heos vespera, S. Kansu, China 

Fig. 

60. 


C. romanovi, Turkestan 

Fig. 

61. 


Anteos menippe, Mattogrosso 

Fig. 

62. 


A. clorinde, Valles, Mex. 

Fig. 

63. 


Catopsilia florella, Syria 

Fig. 

64. 


C. grandidieri, Madagascar 


i(2):t35-i56, 196 } COLIAS GENITALIA 153 

PLATE I 



PETERSEN 


PLATE II 


COLIAS GENITALIA 


PLATE III 





156 


PETERSEN 


/. Res. Lepsd. 


PLATE IV 



6fb\ 


Journal of Research on the Lepidoptera 1 ( 2 ) : 157 - 162 , 1963 

1140 W'. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 1965 

P 


THE EFFECT OF HYBRIDIZATION OF HOST-PLANT 
STRAINS ON GROWTH RATE AND 
MORTALITY OF PIERIS RAPAE' 

WILLIAM HOVANITZ and VINCENT C. S. CHANG 

California Arboretum Foundation, Inc., Arcadia, 

Los Angeles State College and 
University of California, Riverside 


It has been previously shown (Hovanitz and Chang, 1962) 
that larvae of strains of Rieris rapae differ in their mortality rate, growth 
rate and final size dependent upon the plant which is supplied them for 
food. Slightly greater viability, greater rate of growth and increased 
size are characteristic results of feeding the larvae on kale (Brassica 
oleracea var. acephala) as compared with mustard (Brassica nigra), 
even though both plants are utilized by the species in the wild. 

It is the purpose of this paper to report on the results of an experi- 
ment designed to compare these physiological factors in strains bred 
for several generations on each plant 'and then hybridized. 

THE STRAINS 

The strains of Rieris rapae used in these experiments were derived 
originally from two sources, both in the Los Angeles Basin of southern 
California. The first, here designated the kale strain, originated with 
wild females obtained in a cabbage field (Brassica oleracea var. capitata) 
in a truck crop growing area in western Orange County, near Hunting- 
ton Beach. The other, here designated as the mustard strain, originated 
in the fields of the Los Angeles State and County Arboretum, where in 
the spring time Rieris rapae may be found in conjunction with black 
mustard (Brassica nigra). These have probably had no recent contact 
with cabbage since there are none grown commercially within nine 
miles of the area. Before testing in the experiments here described, the 
kale strain had passed through more than ten generations in the labora- 
tory on kale and the mustard strain had passed through more than six 
generations in the laboratory on mustard. 

'Aided by a grant from the National Science Foundation, Washington, D. C. 


157 


158 


HOVANITZ AND CHANG 


/. Res. Lepid. 


THE COMPARATIVE TESTS 

Six comparative tests were run contemporaneously, consisting of 
twenty larvae each. These larvae were fed in petri dishes by daily 
changes of food, keeping each larvae well isolated from the others. It 
was especially important while they were small that the larvae were 
placed directly on top of the leaf of the plant. Of the six tests, two 
were of the parent strains; one of these was derived from kale and fed 
on kale during the experiment, while the other was derived from 
mustard and fed on mustard during the experiment. Four further sets 
were Fi larvae from these two parent strains derived as follows: 

One set consisted of Fi larvae obtained by crossing a kale-strain 
female with a mustard-strain male and then by feeding the ensuing 
larvae on the food-plant of the mother ( kale ) . 

A second set consisted of larvae obtained by crossing a mustard- 
strain female with a kale-strain male and then feeding the ensuing Fi 
larvae on the food plant of the mother ( mustard ) . 

A third set consisted of larvae obtained as in the manner of the 
first F 1 set above, but fed on the food plant of the father ( mustard ) . 

A fourth set consisted of larvae obtained in the manner of the second 
F 1 set above, but fed on the food plant of the father ( kale ) . 


MORTAUTY 

The mortality figures for the larvae were low in all the strains 
( table 1 ) . The kale-strain parents had a mortality of ten percent, which 
compared well with the data previously obtained ( Hovanitz and 
Chang, 1962). The mustard-strain parents had a mortality of thirty 
percent which also compared well with the previously recorded data. 

The reciprocal Fi’s behaved differently. When the kale-strain con- 
tributed the female parent and the mustard-strain contributed the male 
parent, the mortality of the Fi larvae was twenty percent whether kale 
or mustard was used as food. On the other hand, when the mustard- 
strain contributed the female parent and the kale-strain contributed the 
male parent, the larvae had only a ten percent mortality when used on 
kale and zero percent mortality when bred on mustard. It is doubtful 
that these slight differences are significant for the N value of 20, though 
they may represent a real difference. It is possible that the hybrids show 
a greater viability as a result of their heterozygosity. Both parental 
strains have been considerably inbred during their maintenance in the 
laboratory. No reason can be given at this moment for the greater 
viability of the mustard female F/s (10 and 0 percent) as compared 
with the reciprocal kale female F/s (20 percent each). 


i(2):t5y-i62, 196 } 


PIERIS HYBRIDIZATION 


159 


LENGTH OF LARVAL PERIOD 

The minimum length of life for the larvae from the egg to the 
adult for both of the parental strains was twenty-five days under the 
conditions of the experiment (table 1) . All four of the hybrid F, strains 
had a shorter larval length of life indicating a greater activity than the 
inbred parental strains. The rate was speeded up to twenty-two days for 
the Fi strains in which the female parent was derived from kale stock. 


Type 

Strain 

No. 

No. 

No. 

and % 

Minimum days 
from egg to pupae 


Kale-bred 

5 

20 

2 

10% 

25 

Parents 

Mustard-bred 

6 

20 

6 

30% 

25 

u 

Kale-bred 

2 

20 

4 

20% 

22 

Kale ^ X Mustard ^ 

Mustard-bred 

1 

20 

4 

20% 

22 

u 

Kale-bred 

4 

20 

2 

10% 

21 

Mustard ^ x Kale 6^ 

Mustard-bred 

3 

20 

0 

0 

23 


Table 1, The comparative mortality and length of larval growth 


period of various strains of Pieris rapae . 

For the Fi stock in which the female parent was derived from mustard, 
the larvae bred on kale were speeded up to twenty-one days and those 
bred on mustard to twenty-three days. These data definitely indicate a 
significantly greater activity of the hybrid strains than the parental. 
However, there is no indication of any maternal effect on the inheritance 
of food-plant preference since the reciprocal crosses indicated the same 
general effects on mortality and growth rate whichever strain con- 
tributed the female parent. 

GROWTH RATE AND MAXIMUM SIZE 

The growth rate of the six sets of larvae was compared by measur- 
ing ten of the living larvae daily from the time of hatching from the 
egg to the time of pupation. The average of these ten measurements 
was plotted daily and curves drawn to illustrate their size increases 
(fig. 1). Differentiation between the curves occurred within the first 
few days but the first five days have been omitted in fig. 1, owing to 
the small size of the larvae. On the fifth day, both parental strains had 
the smallest larvae while the two F, strains, in which the kale-strain 
female parent was used, had the largest larvae. The other two Fi strains, 
in which the mustard- strain female parent was used, were intermediate. 
During the entire growth period, nearly the same relationships held^ 
with the exception of the kale-bred Fi with a mustard-strain female 


160 


HOVANITZ AND CHANG 


/. Res. Lepid. 



Fig. 1. Growth rates of larvae of six strains of Pieris rapae on kale and 
mustard. Vertical scale, length in mm.; horizontal scale, days after 
hatching from egg to pupation. 


(2):i5j~i62f 1^63 


PIERIS HYBRIDIZATION 


161 


parent which for a time was faster than aM the others. Of the two 
parental groups, the mustard -bred larvae were larger than the kale-bred 
at five days, but the kale-bred reached a maximum ahead of and larger 
than the mustard-bred. Of the three groups on kale as compared with 
the three on mustard, the kale-bred group in each case reached a maxi- 
mum size larger than or earlier than the comparable group on mustard. 
Of the parental as compared with the Fi groups, the larvae of both 
F] hybrid groups consistently exceeded the larvae of the parental 
groups from the beginning to the end. 

The largest maximum size was attained by the F\ group in which 
the female parent was derived from a kale-strain and in which the 
larvae were kale-bred. On the other hand, the second largest maximum 
size was attained by the larvae of the F] group bred on mustard but 
which had a mustard-strain mother. 

It may be observed from the curves that at the periods of molting, 
sizes of the larvae are reduced. These periods are shown by the arrows 
(fig. 1). Because these reduction periods are delayed for the slower 
larvae, the older the larvae, the more difficult it is to compare the six 
groups on any one day. 

SUMMARY AND CONCLUSIONS 

1. Crosses were made between strains bred for many generations 
on kale and mustard. 

2. The effect of hybridization between these host-selection strains 
has been to speed up development, to bring about increased sizes to 
the larvae, and to reduce mortality. These effects are probably due to 
the genetic effects of "hybrid vigor”, in which the deleterious effects of 
inbreeding within the host-selected strains have been reduced by out- 
crossing. 

3 . These effects are greatest for the kale-strain than for the mustard- 
strain. This would tend to confirm our previous findings that- kale is 
preferred as a food over mustard, even when mustard is utilized as the 
food for many generations. 

4. The increased viability, increased growth rate and increased size 
of the larvae are greatest when the hybrid larvae are grown on the plant 
corresponding to the strain of the mother, rather than the father. For 
example, the F-i hybrids where the mother was of the kale-strain grow 
larger and pupate sooner on kale than on mustard. Likewise, the Fi 
hybrids where the mother was of the mustard-strain grow larger but 
not faster on mustard than on kale. 

LITERATURE CITED 

HOVANITZ, W. aod VINCENT C. S. CHANG. 1962. The effect of vari- 
ous food plants on survival and growth rates of Fieru J. Res. Lepid., 
1(1): 21-42. 


162 


HOVANITZ AND CHANG 


/. Res. LepiJ. 


NOTICE 

THE JOURNAL OF RESEARCH ON THE LEPIDOPTERA 
started publication in August, 1962. This fact makes it impossible for 
volume 1 to coincide with the calendar year, 1962 and still have four 
issues to the volume. The editor was confronted with tbe problem of 
two alternatives, ( 1 ) to extend the period for volume 1 to include the 
year 1963 and thus to have only four issues for the year 1962 and 1963, 
or (2) to have volume 1 completed early in 1963 by rapid issuance of 
the remaining numbers and then to issue the four numbers of volume 2 
before the end of the year 1963. The second alternative has been selected, 
and therefore two more issues of the JOURNAL volume 1 will appear 
shortly, to be followed by four issues of volume 2 before the end of 
the year. 

Orders received for the calendar year 1963 with no volume designa- 
tion will be filled with volume 2 only; if volume 1 is desired, the year 
1962 (volume 1) must be ordered. It is hoped that this explanation will 
help to solve the problems of institutional orders which have appeared. 

Orders for individuals who subscribe before the fourth number of 
volume 1 appears, will have their subscriptions start with the first volume, 
and will be billed for volume 2 at the time of sending number 4 of 
volume 1. 

In view of this rapid publication of papers, it is very likely that the 
editor will run short of manuscripts. It is urged that those who have 
good work to publish on the Lepidoptera will take advantage of this 
"chance of the lifetime” and send their MS to the editor. 

It is suggested that subscribers and readers help the JOURNAL 
by becoming a member of THE LEPIDOPTERA FOUNDATION. 
See inside front cover. 


Journal of Research on the ^epidoptera 1(2) ; 163- 168, 1963 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 


CHANGE OF FOOD PLANT PREFERENCE BY 
LARVAE OF PIERIS RAPAE CONTROLLED 
BY STRAIN SELECTION, AND THE 
INHERITANCE OF THIS TRAIT’ 

WILLIAM HOVANITZ and VINCENT C. S. CHANG 

California Arboretum Foundation, Inc. 

Los Angeles State College, Los Angeles 
and 

University of California, Riverside 

It has been shown that larvae which have previously been fed on 
a particular food plant are more likely to select that kind of plant than 
if they had not previously been fed on it (Hovanitz and Chang, 1962 a 
and b). These data have shown that there was a shift in selection of 
mustard or of kale according to whether or not the larvae were from a 
kale or a mustard strain. Likewise, there was shown a greater selection 
for nasturtium by larvae previously fed on nasturtium, and indeed, by 
these same larvae, a greater selection for mustard over kale. This change 
in selection operates whether the larvae were fed on the changed food- 
plant for a long time (several generations) or a short time (part of the 
individual larval life) . 

In this paper, it is our purpose to test the effect of continued 
feeding of the larvae of Pieris rapae for many generations on mustard 
and kale by the process of isolating the strains so fed for many genera- 
tions, then crossing the parental strains to get Fi individuals and finally 
crossing the Fi to get the Fq segregation. 

THE STRAINS 

The strains of Pieris rapae used in these experiments were derived 
originally from two sources, both in the Los Angeles Basin of southern 
California. The first, here designated the kale strain, originated with 
wild females obtained in a cabbage field (Brassica oleracea var. capitata) 
in a truck crop growing area in western Orange County, near Hunting- 
ton Beach. The other, here designated as the mustard strain, originated 
in the fields of the Los Angeles State and County Arboretum, where in 
the spring time Pieris rapae may be found in conjunction with black 
mustard (Brassica nigra). These have probably had no recent contact 
with cabbage since there are none grown commercially within nine 
miles of the area. Before testing in the experiments here described, the 
kale strain had passed through more than ten generations in the labora- 

1 Aided by a grant from the National Science Foundation, Washington, D. C. 


163 


164 


HOVANITZ AND CHANG 


/. Res. Lepid, 


tory on kale and the mustard strain had passed through more than six 
generations in the laboratory on mustard. 

Larvae were selected from each and tested for their seleaion for a 
series of plants in the manner indicated in a previous paper (Hovanitz 
and Chang, 1962). The plants used in these tests were mustard 
(Brassica nigra), kale (Brassica oleracea var. acephala), nasmrtium 
(Tropaeolum majus), Isomeris (Isomeris arborea) and Cleome (Cleome 
huea). 

STRAIN SELECTIONS 

The tests were physically carried out in the manner indicated above. 
In order to be precise on the nature of any difference to be detected by 
these experiments, the tests were conducted in large numbers. Twenty- 
five larvae of each strain were used for twenty or forty trials each, giving 
a total of six hundred test times for each strain (table I). This was 
increased to six hundred sixty for the Fi and nine hundred (with 45 
individuals) for the F 2 . 



mustard 


kale 

nasturtium 

Isomeris 

Cleome 


none 

no. of 

total 














larvae 

test 

PARENTS 















On mustard 

364 

60.66% 

119 

19.83% 

56 

9.33% 

18 

3.0% 

12 

2.0% 

31 

5.16% 

25 

600 

On kale 

144 

24.00% 

354 

59.0 % 

50 

8.33% 

21 

3.5% 

15 

2.5% 

16 

2.66% 

25 

600 

T 

M 9x K cf 

356 

59.93% 

141 

21.36% 

90 

13.63% 

23 

3.48% 

39 

5.91% 

11 

1.67% 

25 

660 

(on mustard) 

9 d- 

(on kale) 

328 

49.68% 

127 

19.24% 

109 

16.51% 

33 

5.00% 

36 

5.45% 

27 

4.09% 

25 

660 

^2 















m'9x K d- 

491 

54.55% 

145 

16.11% 

190 

21.4 % 

30 

3.33% 

42 

4.66% 

2 

0.22% 

45 

900 

(on mustard) 















K9 X M cf 
(on kale) 

406 

45.11% 

146 

16.22% 

219 

24.33% 

79 

8.77% 

39 

4.33% 

11 

1.22% 

45 

900 


Table 1. The comparative mortality and length of larval growth period of 
various strains of Pieris rapae. 

The differential selection of the plants by larvae of the two selected 
strains is clearly made apparent by the curve showing the percentage 
selection (fig. 1). The larvae from the mustard bred strain selected 
mustard sixty-one percent of the time as compared with kale twenty 
percent of the time. The kale strain larvae selected kale fifty-nine 
percent of the time as compared with mustard twenty-four percent of 
the time. The two strains show an almost complete reversal of their 


i( 2 ): i 6 j-i 6 S, 1963 


PIERIS STRAIN SELECTION 


165 


preferences with regard to these two plants, their preference being in 
the direction of the plant utilized as food for several generations. 

The selection by larvae of these two strains for the other plants 
concerned was not significantly different. Nasturtium was selected nine 
percent of the time by the mustard strain and eight percent of the time 
by the kale strain. Isomeris was selected three percent of the time by 
each strain, and Cleome about two percent of the time. About five 



Fig. 1. Curves showing the percentage selection of various food plants by 

larvae of Pieris rapae from a kale and a mustard strain. 

percent of the larvae of the mustard strain left test area with no 
selection as did about three percent of those of the kale strain. Thus, 
there is no difference observable between these two parental strains in 
their selection, except with regard to mustard and kale. 

THE CROSSES 

Adults of the two strains were crossed and the larvae tested for 
their preference toward a selection of plants. Since there was a possi- 
bility of a maternal influence on the inheritance of the food plant 
selectivity hinted at in a previous experiment, reciprocal crosses were 


166 


HOVANITZ AND CHANG 


/. Res. Lepid. 



Fig. 2. Curves of the percentage selection of various food plants by larvae of 
Pieris rapae comparing the reciprocal F, of the strains shown in figure 1. 

made, mustard strain female X kale strain male and vice versa. In each 
case, however, the larvae were bred on the plant of the female strain 
until tested. It is now known that it would also have been desirable to 
have had the reciprocal feeding tests made in addition, as can be seen 
below. 

The F, larvae obtained, in which the female parent was from the 
. mustard strain, showed little difference in selection of plants from the 
parent mustard strain except in an increase in selection of nasturtium 
and Cleome, neither plant of which was involved in this selection. The 
reciprocal Fi larvae obtained from the cross of a kale strain female with 
a mustard strain male showed a much greater selection of mustard 
than the kale parent strain, and was more like that of either the 
mustard parent or of the reciprocal strain indicated (table 1, and fig. 2). 
In fact, the results of this cross would indicate that the genes for 
mustard selection are nearly completely dominant over those for kale 
selection. The cross would also indicate that they are transmitted at 
least through the male since a mustard strain male was used with a kale 
strain female and the larvae were bred on kale. The fact that the larvae 
were bred on kale may be the only reason that the selection was not 
higher toward mustard than is indicated. As with the reciprocal cross, 
there was a great increase in selection of nasturtium and Cleome 
following crossing. This was followed slightly by an increase toward 
Isomeris though this may not be significant. There is little doubt from 
these data that the selection of food plants is inherited, and that 


i(2):i6}-i6S, 196} 


PIERIS STRAIN SELECTION 


167 


"mustard” is dominant over "kale” despite the fact that there is also 
superimposed upon this inheritance a selective propensity controlled 
by training during the life of the individual, as has been shown before. 

The F 2 crosses were made in the same manner. F 2 larvae of each of 
the crosses indicated above were obtained and bred on the food plant 
of the mother ( table 1 ) . The tests on these larvae led to basically the 
same results as on the Fi larvae with the exception that all percentages 
were reduced slightly and the percentages for nasturtium were raised 
greatly. Of significance too is the reduction of the larvae which made 
no selection at all. The reduction in selection of mustard and kale 
appears proportional and related to the increase in the selection of 
nasturtium (fig. 3 ) . 

DISCUSSION AND CONCLUSIONS 

The data indicated in this paper appear to show that selection of 
genes for food plant preference occurs over a period of generations 
when strains are maintained isolated on particular food plants for many 
generations. These genes do not show maternal inheritance. In the 
crosses indicated in this paper, genes for selection of mustard over kale 
are apparently dominant over the reverse. This is indicated by the fact 
that in the Fi of the cross kale strain X mustard strain or the reciprocal, 
selection is similar to that shown by the original mustard strain. The 
same results are shown for the F^ but they are not so pronounced. Of 
great curiosity is the fact that in the Fi there is an increase in selection 
for nasturtium, not one of the preferred plants of Pieris rapae ( Flovanitz 
and Chang, 1962a). Feeding larvae on nasturtium has shown that there 
may be greater selection of this plant developed than would ordinarily 
be present. In the present case, however, no selection of this sort is 
involved and the increase is significant. This increase is even greater 
in the F 2 cross than in the Fi, there being a two and one half- to three- 
fold increase in selection of nasturtium in the F 2 as compared with the 
parental strains. A much slighter increase is also indicated for Isomeris 
and Cleome, and a decrease for the number of rejects, that is, those that 
make no selection. Reasons for this much greater selection for nasturtium 
and other plants following hybridization are not known nor can they 
even be guessed at intelligently at this time. 

SUMMARY 

1. Two food plant strains were developed by selection, one on 
kale (Bras sic a oleracea) and another on mustard (Brassica nigra). 

2. When tested, each of these strains showed a much greater 
preference for their accustomed food plant than for any other tested. 
Those tested were mustard, kale, nasturtium, Isomeris and Cleome. 

3 . When these two strains are crossed, the Fi hybrids showed a 
preferential selection most like the mustard parent strain. This indicates 
that the gene(s) for mustard is dominant over that for kale. There is 


168 


HOVANITZ AND CHANG 


J. Res. Lepid. 



Fig. 3. Curves of the percentage selection of various food plants by larvae of 
Pieris rapae comparing the reciprocal F 2 of the strains shown in 
figures land 2. 

also indicated a slight increase in selection of the other plants tested. 

4. The Fo of this cross showed results similar to the Fi, namely a 
selection in favor of mustard rather than kale whether or not the female 
parent had originally come from the mustard or the kale strain. Thus, 
maternal inheritance is not indicated here even though maternal effects 
were indicated on growth rate and mortality data in a previous paper. 

5. As had been indicated, feeding the larvae even a short time on 
one particular food plant may influence it to have a preferential 
selection for that plant. 

6 . In the F2 there is indicated a strongly increased preferential 
selection toward nasturtium which was not selected in any previous 
strain. This increase is also indicated slightly toward Isomeris and 
Cleome. No reason is advanced at this time for these results. 

LITERATURE CITED 

HOVANITZ, W. and VINCENT C. S. CHANG. 1962a. The effect of vari- 
ous food plants on survival and growth rates of Pieris. J. Res. Lepid. 
1(1): 21-42. 

1962b. The effect of hybridization of host-plant strains on 

growth rate and mortality of Pieris rapae. J. Res. Lepid. 1(2) : this issue. 


advertisement 


TAKi CHLOROCRESOL ON YOUR NEXT TRIP 


Since the publication of the Chlorocresol method, 
field collecting techniques have been changed radi- 
cally. The Chlorocresol method eliminates the need 
for time consuming field mounting and/or relaxing 
of specimens. With Chlorocresol, specimens are re- 
tained in a relaxed condition for an unlimited period, 
allowing mounting to be done at a more convenient 
time or place. Field mounted specimens which would 
normally occupy a standard insect box may be packed 
in a container only 5" x 5" x IV 2 '. With specimens 
in a relaxed state, damage is minimized in the event 


of rough handling. 

Catalog No. 182 

Chlorocresol, 50 grams $0.95 

Chlorocresol, 100 grams 1.75 

Chlorocresol, 200 grams 3 00 

Styrene box, 5" x 5" x IV 2 ' per dozen 3.00 


Complete instructions are included with each 
shipment. 

Prices are F.O.B. Santa Monica, California. California 
residents please add 4% sales tax. 

General Entomological Supplies 



BIO METAL ASSOCIATES 

Box 61 

Santa Monica, California 


Volume 1 


Number 2 


January, 1963 


THE J@U^NJAL ©F RESEARCH 
©NJ THE LEFIJ©©RTERA\ 


IN THIS ISSUE 

Composition and relative abundance in a temperate zone 
butterfly fauna . . . Thomas C. Emmel and John F. Emmel 97 

The Argynnis populations of the Sand Creek area, Klamath 
Co., Oregon, Part I J- W. Tilden 109 

Caterpillar Versus Dinosaur? . . . Theodore H. Eaton, Jr. 114 

Geographical distribution and variation of the Genus Argynnis 
I. Introduction 

II. Argynnis idalia William Hovanitz 117 

The relation of Pieris virginiensis Edw. to Pieris napi L. 

Species formation in Pieris? Wiliam Hovanitz 124 

The male genitalia of some Co lias species . Bjorn. Petersen 135 

The effect of hybridization of host -plant strains on growth rate 

and mortality of Pieris rapae 

William Hovanitz and Vincent C. S. Chang 157 

Notice-editorial 162 

Change of food plant preference by larvae of Pieris rapae 
controlled by strain selection, and the inheritance of this trait 

William Hovanitz and Vincent C. S. Chang 163 



@F 

OIMJ THE LEFI1P@PTER1A 


MeSfeii 


Bi2!!*S*****»»*«'*" . 

... 

^^■rs****** ***•*•» 




!psr.i:j2:r*fe; 

us:sss:2s;*.2ui,. 

K«sjssK2.»:2i;ir 


Volume 1 


Number 3 


March, 1963 

i, rr V 


* 




Established in 1 962 
Published at 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

Edited by WILLIAM HOVANITZ 

WITH EMPHASIS ON ENVIRONMENTALLY AND GENE- 
TICALLY INDUCED VARIATION, population analysis, evolution, 
phylogenetic taxonomy, zoogeography, comparative morphology, ecol- 
ogy, geographical variation, speciation, physiology, etc. In short, 
quality luork on any aspect of research on the Lepidoptera. 

THE PURPOSE OF THE JOURNAL is to combine in one source 
the work in this field for the aid of students of this group of insects 
in a way not at present available. The JOURNAL will attempt to 
publish primarily only critical and complete papers of an analytical 
nature, though there will be a limited section devoted to shorter 
papers and notes. 

RATES: $ 8.00 per volume, personal subscription. 

$12.00 per volume, institutional subscription. 

All amounts are in U.S. dollars, payable in the U.S.A. 

AUTHORS ARE REQUESTED to refer to the Journal as an 
example of the form to be used in preparing their manuscripts. Fifty 
separates will be supplied to authors free; reprints will be sold at 
printer’s rates if ordered at time galley proofs are returned. If proofs 
are not returned promptly, the editor reserves the right to withhold 
publication, or to proceed with no responsibility. All issues of the 
Journal will be copyrighted; in submitting a paper each author agrees 
that the material has not been published elsewhere. 

MANUSCRIPTS AND SUBSCRIPTIONS should be mailed to the 
address noted above. 

THE LEPIDOPTERA FOUNDATION 

To provide permanent security to the JOURNAL, THE LEPIDOP- 
TERA FOUNDATION is being set up. Memberships are now being 
accepted at TEN DOLLARS per year, eight dollars of which goes to 
your subscription and two dollars goes into a permanent capital fund, 
the earnings only of which may be used for current publishing expenses. 
In addition a scale of higher contributions is to be set up which will 
be announced later. All funds received in excess of the subscription rate 
will go into the permanent fund. LIFE MEMBERSHIPS are offered 
now at Two hundred fifty dollars; these include the Journal for life. 
INSTITUTIONAL AND COMMERCIAL DONATIONS AND MEM- 
BERSHIPS are also available; rates will be announced later, or on request. 
These contributions are deductible. HELP YOUR JOURNAL. Meetings 
of members may be held at times and places to be arranged. 


Journal of Research on the Lepidoptera 1 ( 3 ) :169-182, 1963 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
© Copyright 196 } 


SELECTION OF ALLYL ISOTHIOCYANATE BY 
LARVAE OF PIERIS RAPAE AND THE 
INHERITANCE OF THIS TRAIT' 


WILLLIAM HOVANITZ and VINCENT C. S. CHANG 

California Arboretum Foundation, Inc., Arcadia, 

Los Angeles State College and 
University of California, Riverside 

Larvae which have previously been fed on a particular food 
plant are more likely to select that plant than if they had not previously 
been fed on it (Hovanitz and Chang, \9G2a and ^). It was shown that 
larvae which were fed on kale are more likely to select kale, and larvae 
which were fed on mustard are more likely to select mustard. Also, 
larvae which have previously been fed nasturtium have a higher 
selection on that plant than if they had not been fed on it. This 
change in food plant selection operates both if the adaptation comes 
over a period of several generations, or if only for a short time in the 
individual larval life. Later, it was shown that strains selected for 
several generations on each of two plants, kale and mustard, had strong 
selection for their respective food plants ( Hovanitz and Chang, 
1963 ^^ and b). Individuals from these two strains were then crossed to 
get the Fi generation, and these were crossed to get the Fo generation. 
The inheritance of this trait appears to be multi factorial but because 
of the nature of the testing, this point may be open to question. 

The data to be presented in this paper were obtained in much the 
same manner as in the previously mentioned papers (Hovanitz and 
Chang, 1963<? and b) with the exception that instead of testing the 
selection of the larvae for the plants themselves, they were tested for 
their selection of various concentrations of black mustard oil (allyl 
isothiocyaoate ) . Tests were made on the parental larvae as well as the 
Ft and the F 2 . 

THE STRAINS 

The strains of Pieris rapae used in these experiments were derived 
from two original sources. The first, the kale strain, originated with wild 
females obtained in a cabbage field {Brassica oleracea var. capitata) 
in a truck crop growing area near Huntington Beach in Orange County, 
California. The other, the mustard strain, originated in the fields of the 


'Aided by a grant from the National Science Foundation. 

169 SlllTflSIJNlW 

mSTlTUTiaH 


APR1819® 


170 


HOVANITZ AND CHANG 


/. Res, Lepid. 


Los Angeles State and County Arboretum where P. rapae is found on 
black mustard (Brassica nigra). Before testing in the experiments here 
described, the kale strain had passed through more than ten generations 
in the laboratory on kale {Brassica oleracea var. acephala) and the 
mustard strain had passed through more than six generations in the 
laboratory on black mustard. 

EXPERIMENTAL SET-UP 

The tests of the larvae were made in much the same manner as has 
previously been described (Hovanitz and Chang, 1962b) except that 
in place of the plants to be selected, at various points surrounding the 
flat, filter papers wetted with various solutions of allyl-isothiocyanate 
were placed. Preliminary tests had indicated that the larvae were 
attracted to dilutions of the mustard oil at 10"^ to 10"'^ so the tests were 
made at these dilutions. Metal mosquito netting was cut in a circular 
arrangement to serve as a base for the filter paper. These discs of metal 
netting were placed on top of glass flasks partially buried in soil, secured 
to corks placed in the flasks and arranged in various parts of the flat 
according to the disposition shown in Figure 1. The filter paper was 
then set up top of the metal netting and wetted with distilled water 
solutions as shown. On one of these, distilled water only was placed so 
as to serve as a control for the others. To prevent the accumulation of 
mustard oil concentrations, the solutions were alternated around the 
flat. The set-up of the experiments is shown diagrammatically in Figure 1 
and photographically in Figure 2. The larvae were set in the center of 
the flat and allowed to select any of the six test papers or to leave the 
flat without selection. It should he noted that the test papers were above 
the level of the larvae and therefore that the heavier-than-air mustard 
oils would drop down to their level. After each trial, the soil in the flat 
was mixed to prevent the origin of "trails.” It should also be noted that 
it would be much easier for the larvae to leave the flat without any 
selection except for one factor, namely, that these larvae have a tendency 
to climb upwards when at all possible. Experience with lepidopterous 
larvae has shown that descent is usually made by means of dropping 
down on a silken thread, rather than by walking while ascent is made 
by walking. This accounts for the selection of the "water” alone just 
as frequently as they leave the flat without any selection at all. 

The mustard oil used in these experiments was a natural product 
obtained from black mustard seed by compression and distillation. It 
is available commercially from the Fritsche Brothers firm. Allyl iso- 
thiocyanate is the principal component of the oil obtained from Brassica 
nigra; other mustard oils are found in other plants and all are esters of 
isothyocyanic acid. Brassica nigra seeds contain a glucoside, potassium 
myronate or sinigrin, which undergoes fermentation change due to the 
enzyme myrosin present in the seeds. 


(})ti69-ii2, 196} 


ALLYL ISOTHYOCYANATE 


171 



Fig. L The set-up for the larval tests. The larvae are placed in the 
center of the flat from where they are enabled to go in any direction toward 
one of the dilutions of mustard oil, or to leave the flat. The circles show the 
disposition of the filter papers which have upon them dilutions of mustard 
oil as indicated. 

THE TEST SELECTIONS 

Test selections were made by using a number of larvae for several 
times to get a combined sample. The number of individuals used 
depended in part on those available, but ranged from fifteen to forty-five 
giving a total number of tests from three hundred to nine hundred. 
These data are given in Table 1. 

The parent strains were tested first. The mustard strain was tested 
with twenty -five larvae 20 or 40 times each giving a total of seven 
hundred trials. Of these, only nineteen or 2.7 percent made no selection 
and left the flat. Only twenty-four or 3.4 percent selected the filter 
paper with distilled water only on it. The others all selected the mustard 
oils, from a low of nearly 10 percent for the 10"’ to a high of nearly 
thirty percent for the 10"^. These data are shown diagrammatically in 
Figure 3 . 


172 


HOVANITZ AND CHANG 


/. Kes. Lepid. 



Fig. 2: Photograph showing the surface of the experimental flat, the 
flasks with filter paper upon them and a larva raising its foreparts. The rules 
are not present during an experiment and are indicated only for the purpose 
of illustrating the dimensions between the soil surface, the filter paper and 
the larvae. 


for different concentrations of allyl isothiothiocyanate 

(black mustard oil). 


1(3)^169-182, 1963 


ALLYL ISOTHIOCYANATE 


173 


5 



174 


HOVANITZ AND CHANG 


/. Res. Lepid, 


The kale strain was also tested with twenty-five larvae for a total 
of five hundred eighty trials. The highest selection of these larvae was 
toward the concentration of 10"®, larvae going to this concentration in 
thirty-four percent of the trials. This result is significantly different 
from the result for the kale larvae indicated above, in which the highest 
selection was toward the 10"® concentration. These data can be com- 
pared diagrammatically in Figure 3. There is little doubt that the larvae 
of the kale-bred stock prefer the mustard oil at a lower concentration 
than do those of the mustard strain. It is probably not coincidental that 
the taste of the leaves of mustard is stronger than is that of kale (or 
cabbage) since the significant taste of these plants is due to allyl or 
other isothiocyanates. These data would seem to indicate ( 1 ) that the 
concentration of taste or smell-detectable mustard oils in the leaves 
of mustard is higher than that in kale, (2) that the larvae feeding on 
mustard have become adapted in some way to a higher concentration 
of these oils, and (3) that due to this adaptation, the "mustard” larvae 
have a greater selection of the higher concentration of mustard oil than 
the lower. It has been shown in another paper (Hovanitz and Chang, 
1963b) that these two strains also select their own plants over the 
others when given a free choice. This preferred selection may be due 
wholly or in part to the difference in available allyl or other isothiocya- 
nate in their leaves. 


THE CROSSES 

The crosses between the two food plant strains were carried out by 
making reciprocal hybrids. In the one case, the kale strain was used as 
the female parent and in the other case, the mustard strain. The hybrid 
larvae resulting from these crosses were fed on the food plant of the 
mother, since the mother was allowed to lay on its own preferred plant. 

The selections made by the hybrid larvae in both crosses favored 
the concentration 10"^, exactly half way between the selections made 
by the respective parent strains ( Fig. 4 ) . The frequency of selections 
for the kale female Fi toward the 10"^ concentration was 34 percent as 
compared with 30 percent for the mustard female Fi. 

Previous data have indicated that the immediate larval food plant 
eaten by the larvae had an effect on the food plant selections made by 
these larvae. If this held true for the selections of the mustard oils also, 
then the larvae of the kale female Fi, having been fed on kale during 
their life, and the larvae of the mustard female Fi, having been fed 
on mustard during their life, should have a preference for their respec- 
tive plant. Data from Table 1 (Hovanitz and Chang, 1963b) indicate 
that this is so. Mustard female Fi larvae selected mustard nearly 60 
percent of the total selections as compared with nearly 50 percent selec- 
tions for the kale female Fi larval selections. It should follow, if our 


PERCENT SELECTION PERCENT SELECTION 


i(3):i69-az, 196) 


ALLYL ISOTHIOCYANATE 


175 




Fig. 3: Histograms illustrating the selections made by the larvae from 
the parental strains, comparing the kale and the mustard strains. 


PERCENT SELECTION PERCENT SELECTION 


176 


HOVANITZ AND CHANG 


/. Kes. Lepid. 




Fig. 4: Histograms illustrating the selections made by the Fi 
from the reciprocal crosses between the kale and mustard strains. 


larvae 


PEHCEMT SELECTiON PERCENT SELECTION 


(3);i6^-ii2, 1^63 


ALLYL ISOTHIOCYANATE 


177 




Fig. 5. Histograms illustrating the selections made by the Fa 
from the reciprocal crosses between the kale and the mustard strains. 


larvae 


178 


HOVANITZ AND CHANG 


/. Res. Lcpid. 


assumptions are correct concerning the relationship between mustard 
oil concentration and food plant selectivity, that the mustard female Fi 
larvae should select mustard oil concentrations at a higher level than 
the kale female F] larvae. That this is so can be seen by study of the 
histograms (Fig. 4 ) and the following calculations. Selections made 
by the mustard female Fi larvae which are less than 10"^ (that is 
10 "^ or ‘^) are 31.4 percent of the total as compared with 24.6 percent 
of the total for the kale female F, larvae. This shows that the larvae of 
the mustard female Fi select a higher concentration of mustard oil than 
those of the kale female Fi even though the modal class in both cases 
is at 10"^. This difference is most probably attributed to the fact that 
the immediate F, larvae had been fed on the food plant of the mother, 
rather than that there was any genetic effect of maternal inheritance. 

Two sets of Fq have been bred, corresponding to the two sets of Fi. 
These two sets of Fi are the reciprocal crosses mentioned above. The Fq 
from the mustard female Fi was continued on mustard for the food 
plant of the F 2 larvae, and the F 2 from the kale female Fi was con- 
tinued on kale. It would be expected, therefore, that each of these 
strains might diverge slightly in their selection according to the pattern 
of the larval foods. This result did not materialize. 

The data indicate that the Fq of both strains still have a selective 
propensity favoring the dilution, 10"^, thus having the same modal 
class as the F 1 ( Fig. 5 ) . This is the expected genetic result for multiple 
factor inheritance. The second result expected for multiple factor in- 
heritance is a wider range of variation in the F 2 than the Fi. This 
result is not shown by a wider range per se since no wider range of 
dilutions were used, but rather it is shown by a greater dispersion of 
the selections inside the range tested. This would give a greater standard 
deviation for the F 2 than for the corresponding Fi. The greater disper- 
sion is very clear by study of the histograms, rearranged so that the 
mustard strains are on one page and the kale strains are on another 
(Figs. 6 and 7). The differences between the Fi and the F 2 are striking. 
These results indicate clearly that the factors for food plant selection in 
Pieris rapae which have been separated in the two strains here tested 
are genetic and behave in a way comparable with polygenic inheritanp. 
It also indicates that the populations of Pieris rapae existing in the wild 
are differentiated into genetically distinct populations, differing by genes 
which control the selection of food plants. These differences could 
account for a mechanism needed for species isolation (Hovanitz 1963c). 

DISCUSSION AND CONCLUSIONS 

The data indicated in this paper show that the strains which have 
been selected for particular food plant preferences over a period of 
several generations, and which when tested prefer the food plant upon 
which they have fed, also differ in their selection (preference) for vari- 




ALLYL ISOTHIOCYANATE 


179 


ous concentrations of mustard oil (allyl isothiocyanate). That these 
differences are genetic is apparent not only from the preceding data 
on food plant preferences but also on the data on mustard oil preferences 
introduced in this paper. Crossing of the two strains discussed here gives 
an F, which has a selection exactly intermediate between the two par- 
ental strains. The results are similar whether the kale strain contributes 
the maternal parent or whether the mustard strain contributes the 
maternal parent. There is, however, a greater selection of the higher 
concentrations in the strain fed on mustard than the one fed on kale. 

The F 2 from these strains is similar to the Fi except for the wider 
dispersion of the selections. 

In the testing of these strains for food plant preferences, it was 
found that the larvae developed a preference for nasturtium in the Fi 
and F 2 . No reason can be deduced for this strange event based upon the 
present results unless it be that nasturtium emits at a concentration of 
mustard oil of the same attractive power as 10"^ allyl isothiocyanate or 
some other mustard oil of attractive significance. The odor of nasturtium 
does not seem to the human to be the same as that of allyl iosthiocyanate. 
However, it is possible that to the human, the odor of some other sub- 
stance masks the odor of the attractive mustard oil, but that this odor 
does not do so in the larvae which are able to detect it. 

Thorsteinsen (1953) has made the statement that larvae of the 
diamond-back moth, Plutella macuUpennis (Cut.) are attracted to 
the odor of ally! isothiocyanate from a distance but are repelled by it 
when closeup so that they will not eat substances having this oil. Instead, 
they will eat substances with the corresponding glucoside, potassium 
myronate, but are not attracted to this substance from a distance. Since 
the glucoside is converted into the allyl isothiocyanate in the plant by 
a process of fermentation, it is probable that the attraction is really for 
the isothiocyanate not the glucoside, but that the isothiocyanate is pro- 
duced slowly and in small quantities by the process of enzymatic 
fermentation. Indeed, Thorsteinson even indicates "in some experi- 
ments the addition of allyl mustard oils slightly increased feeding on 
media containing sinigrin.” Wolfrom (I960) points out that "while 
the exact biological function of the plant glycosides is not established, 
it is probable that their formation provides the plant with a means of 
storing, in a harmless form, toxic and physiologically active materials 
which may be liberated by enzymes, in small quantities, when required.” 
It seems not only possible but highly likely that the attraction differ- 
ences noted by Thorsteinson are based upon the low concentration of 
allyl isothiocyanate rather than upon the presence of the glucoside 
sinigrin directly. Thorsteinson in fact admits this possibility in his com- 
ment, "On the other hand, it is possible that infinitesimal amounts of 
mustard oil vapor emanating from the leaves [by enzymatic action of 
myrosin on sinigrin] may stimulate the olfactory sense which is char- 
acteristically extremely sensitive in insects.” Section in brackets not 


PERCENT SELECTION PERCENT SELECTION 


180 


HOVANITZ AND CHANG 


/. Res. Lepid. 




Fig 6. Histograms illustrating the selections made by the Fi and F:; 
larvae, comparing these larvae of the kale female crosses. 


PERCENT SELECTION PERCENT SELECTION 


(}): i 69- i 82, 196} 


ALLYL ISOTHIOCYANATE 


181 




Fig. 7. Histograms illustrating the selections made by the Fi and F 2 
larvae, comparing these larvae of the mustard female crosses. 


182 


HOVANITZ AND CHANG 


/. Res. Lepid. 


Thorsteinson’s. Further testing of feeding responses of Plutella or 
Pieris larvae at a low mustard oil level are required to be certain of 
this differentiation. Presently published data of the present, or of other, 
authors are definitely not conclusive. Tests of this sort may be very 
difficult to make due to the high volatility of the mustard oil. There is 
no doubt that the use of the glucoside, rather than the oil, makes easier 
the maintenance of a low level of the oil vapor in the vicinity of the 
food. Our own data comparing the egg laying of the adults which are 
to be published later shows that the glucoside and mustard oil released 
from crushing seeds is more attractive than the mustard oil alone. 

Verschaffelt (1910) was the first to study the relationship between 
the food plant choice of insects, plant odor and the chemistry of the 
relationship. He supposed that the larvae were attracted to the mustard 
oils present in the food plants but did not know whether the larvae 
differentiated between different oils or not. Dethier (1941) showed 
the selective effect of various odiferous oils toward lepidopterous larvae. 
Johansson (1951) showed that Pieris hrassicae larvae preferred food 
plants to which they had already become accustomed by previous 
eating. This result has also been shown by us (Hovanitz and Chang 
1962b and 1963a) on Pieris rapae. 


LITERATURE CITED 

DETHIER, V. G. 1941. Chemical factors determining choice of food pianis 
by Papilio larvae. Amer. Nat. 75: 61-75. 

HOVANITZ, WILLIAM and VINCENT C. S. CHANG. 1962a. The effect 
of various food plants on survival and growth rate of Pieris. /, Res. 
Lepid. 1.(1): 21-42. 

— — ■. 1962b. Three factors affecting larval choice of food plant. 

J. Res. Lepid. 1(1): 51-62^ 

— — — ■. 1963a. The effect of hybridization of host-plant strains on 

growth rate and mortality of Pieris rapae. J. Res. Lepid. 1 (2) : 157-161. 

• — . 1963b. Change of food plant preference by larvae of Pieris 

rapae controlled by strain selection, and the inheritance of this trait. 
/. Res. Lepid. 1 (2) ; 163-168. 

. 1963c. The relation of Pieris virginiensis Edw. to 

Pieris napi L. Species formation in Pieris? /. Res. Lepid. 1 (2) ; 124-134. 

JOHANSSON, ARNE SEMB. 1951 The food plant preference of the larvae 

of Pieris hrassicae L. Norsk Ent. Tid. B. VIII, 2.4-5: 187-195. 

THORSTEINSON, A. J. 1953. The chemotactic responses that determine 
host specificity in an oligophagous insect {Plutella maculipennis (Curt.) 
Lepidoptera ) . Canad. J. Zool. 31: 52-72. 

VERSCHAFFELT, E. 1910, The cause determining the selection of food in 
some herbivorous insects. Proc. Acad. Sci. Amsterdam 13(1): 536-542. 

WOLFROM, M. L. I960. Natural glycosides. Encyclopaedia Britannica 
10: 448.. 


Journal of Research on the Lepidoptera 1 (3 ); 183=190, 1963 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 1 ^ 6 } 


BIOLOGY OF THE CEANOTHUS STEM-GALL MOTH, 
PERIPLOCA CEANOTHIELLA (Cosens) 

With Consideration of its Control* 

J. ALEX MUNRO 

Los Angeles State and County Arboretum 
Arcadia, California 

The stem-gall moth, Periploca ceanothiella {Cosens), has 
received but slight attention considering its importance as a pest of 
ornamentals on the genus Ceanothus. The genus Ceanothus, according 
to Van Rensselaer and McMinn (1942), comprises some 55 species 
native only to North America. Of these, some 44 species, and a number 
of horticultural varieties or selections, are grown in the Pacific Coast 
states. Many of them exhibit rather colorful bloom, ranging from the 
white flowers of C. americanus (New Jersey Tea) to the deep blue, 
lilac -like clusters of C. thyrsifloms (Blue Blossom) and the pink of 
some selections. They occur mainly as shrubs and are used in land- 
scape plantings and roadside beautification. Some, such as C. gloriosus 
exaltatus (Point Reyes Creeper) and C. griseus horizontalis (Carmel 
Creeper) are used as attractive groundcovers because of their low 
growing habit. 

The distribution of the gall moth probably corresponds to the range 
of its host. Hodges (1962) in reviewing the genus Periploca mentions 
having examined specimens of P. ceanothiella collected in California, 
Kansas, New York, Ontario (Canada), and Texas. His review shifted 
this insect from the genus Stagmatophora, which it formerly occupied, 
to its present position in the genus Periploca. 


DEVELOPMENT 

This insect normally overwinters within its gall in some stage of 
larval development. Reference to Table 1 shows pupation to be barely 
under way in April and to continue, with moth emergence, throughout 
spring and early summer. Emergence, however, may begin as early as 
January from mature larvae brought in from outdoors and maintained 
under room temperatures. 

The small, dark-colored moths (Fig. 1) begin egg laying within a 
few days following emergence. They deposit their small (0.5 mm) white 

■^Presented at the annual meeting of the Entomological Society of America, Phoenix, Arizona — 
December 3-6, 1962. 


183 


184 


MUNRO 


/. Res. Lcpid. 


TABLE 1. Status of overwintered Periploca ceanothiella galls. 


PLACE 6 DATE 


PERCENT GALLS 

1962 

Number of 
galls exam 

With 

Larvae 

With 

Pupae 

With 

Parasites 

Emerged 

Arcadia 

April 9 

88 

87 

2 

11 

0 

May 18 

64 

60 

21 

15 

4 

June 7 

142 

23 

44 

19 

14 

July 26 

104 

2 

6 

3 

89 

Claremont 

April 25 

85 

52 

28 

18 

2 

May 16 

80 

28 

22 

11 

39 

June 14 

76 

19 

26 

15 

40 

La Canada 

April 17 

66 

94 

0 

6 

0 

May 22 

70 

62 

4 

19 

15 

Santa Barbara 
April 14 

141 

95 

1 

4 

0 


eggs on the underside of leaves, in leaf axils, and along the twigs. The 
eggs required 10 days for hatching under indoor temperatures of about 
70° F. The newly hatched larvae move directly into the terminal growth 
where they penetrate the buds and inflorescences to cause the swellings 
or galls. Here the larva remains, feeding and growing on the inner 
tissue, until reaching full growth of 5 to 6.5 mm by the following 
spring or summer. 

Before pupation begins, the larva cuts an exit hole partially through 
the side of the gall to leave only a thin film of bark which covers the 
entrance until time for the moth to emerge. This "window” or blocked 
entrance provides protection for the occupant against some enemies 
until it is ready to escape from the gall cavity. The pupation stage is 
spent within the gall and requires 24 days under room temperatures of 
about 70 °F. Although the gall moths may emerge over a prolonged 
period of spring and summer, only one brood or annual generation was 
observed. 

The pest has a stunting effect on the new growth. The galls may 
vary greatly in abundance, ranging from only an occasional gall to 20 
or more per lineal foot of branch and twigs in heavy infestations. Some 


(^)-.iS3-i90, 19^3 


GALL MOTH 


185 


of the twigs may be killed outright from the injury. The most serious 
damage, however, was to the bloom. (Fig. 2). It was not unusual to 
see the flower clusters reduced to about 25% of their normal size, 
particularly where the gall larvae were present in the inflorescenses. 

In no instance was more than one larva found per gall. In appear- 
ance, the gall is somewhat spindle-shaped and averaged about 13mm 
in length by 6mm diameter. (Fig. 3) This is about three times the 
diameter of the twig at point of junction with the gall. 

PARASITES 

The role of three naturally occurring ichneumonids in checking 
the stem gall is indicated in Table 1. All three, Fristomems haumhoferi 
Cush., Scambus aplopappi (Ashm.), and Apistephialtes nucicola 
(Cush.), were observed mainly during the spring months. No hyper- 
parasites were observed. Identifications were made by L. M. Walkley 
of the U.S. National Museum Staff. Muesebeck, Krombein, and Townes 
(1951) mention these ichneumonids as parasites of certain micro- 
lepidopterous larvae that tunnel in stems. Of the three parasites, P. 
haumhoferi was the most commonly encountered, and was the only 
one previously recorded as a parasite of this stem gall insect. The toll 
taken by the parasites ranged up to 19% of the gall-larvae. Only the 
mature larvae were attacked by the parasites. 

NON-SUSCEPTIBILIl’Y IN CEANOTHUS 

Examination of Ceanothus collections at the Los Angeles State 
and County Arboretum in Arcadia, Rancho Santa Ana Botanic Garden 
at Claremont, and the Santa Barbara Botanic Garden at Santa Barbara 
(all southern California locations) showed varying degrees of non- 
susceptibility to the stem galls. It was an ideal way to check on this 
factor inasmuch as the Ceanothus species and varieties growing in each 
collection were equally exposed to the pest. The results presented in 
Table 2 would suggest the use of non-susceptible Ceanothus in plant- 
ings wherever practicable. 

CHEMICAL CONTROL 

Inasmuch as spraying of the Ceanothus with DDT had given little 
or no apparent control of this pest it was decided to test the systemic 
insecticide, dimethoate, for the purpose. A commercial formulation 
(Cygon) containing 43.5% dimethoate was mixed at the rate of one 
pint of this concentrate to 100 gallons of water and applied as a foliar 
spray to Ceanothus plantings in Arcadia on June 27, 1962. Examina- 
tion made after one week following application showed that 92% of 


186 


MUNRO 


/. Res. Li pid. 


TABLE 2, Occurrence of Periploca ceanothlella on various species, varieties 
and horticultural selections of Ceanothus. 


HEAVY OCCURRENCE ; 

Ceanothus griseus 
C, griseus horizontalis 


MODERATE OCCURRENCE; 

C. cyaneus 
C, thyrsiflorus 
C, 'Ray Hartman* 
C, 'Marie Simon' 


LIGHT OCCURRENCE: 

C. arboreus 
C, oliganthus 
C, divers if olius 
C. lemmonii 
C, integerrimus 
C, leucodermis 
C, lobbianus 
C, 'Treasure Island* 
C. 'Sierra Blue* 

C, 'Royal Blue* 

C, 'Mountain Haze* 

C, 'Mary Lake' 

C. 'Concha* 


NO OCCURRENCES 

C, americanus 
C, parry i 
C. papillosus 
C, impressus 
C, foliosus 
C. insularis 
C. cuneatus 

C« ramulosus fascicularis 
C, gloriosus 
C. gloriosus exaltatus 
C. rigidus albus 
C. purpureus 
C. prostratus 
C. verrucosus 
C. spinosus 
C, jepsonii 
C, raasonii 
C« megacarpus 
C, greggi perplexus 
C, 'Blue Cloud* 

C. 'Lester Rowntree* 


the gall larvae were killed. The spray, however, had no appa'rent effect 
on the mature gall larvae (which had ceased feeding), the pupae, or 
the parasites, P. baumhoferi and A. nucicola; adults of these normally 
emerged from galls collected from the treated areas. Effect of the treat- 
ment on 5’. aplopappi was not observed. The parasites developed only 
in the mature gaU larvae, and therefore it is unlikely that they had had 
any contact with the insecticide. In addition to checking the stem gall 
infestation, the dimethoate also controlled the Ceanothus leaf miner, 
Nepticula ceanothi Braun, thrips, and an infestation of mites which 
were troublesome at the time. The spray used at this rate had no ob- 
servable phytotoxic effect on the Ceanothus. 

A similar application made on September 14, 1962 showed only 
74% mortality of the gall larvae. This lower kill may have been 
due to the greater proportion of older, possibly more resistant, larvae at 
this time than at the June 27th spraying. 

A soil drench of the dimethoate, made at six times the strength of 
the foliar spray, applied around the base of Ceanothus shrubs in Pasa- 
dena on July 11, 1962, resulted in 70% mortality of the gall larvae, but 


196} 


GALL MOTH 


187 




M E T R I C 


.j_,. 


J_j_j. 


1 1, i 


S Y S T E .M 


Fig. 1. The adult moth, Periploca ceanothiella (Cosens), at bottom 
right; larva and pupa at left; larva in cut gall at top. 


188 


MUNRO 


/. Res. Lepid. 



I> 


i(}):iS}-i90, 196} 


GALL MOTH 


189 



Fig. 3. Stem galls from which the moths have emerged; note the exit 
hole in the encircled gall. The twig at the right has been completely killed by 
the larvae. Note that the galls occur at the beginning of each period of stem 
growth, and that in the twig at the left, two different years’ galls are shown. 


190 


MUNRO 


/. Kcs. Lcpid. 


caused burning and subsequent defoliation wherever the drench came 
in contact with the low-growing foliage. Examination of galls from 
untreated Ceanothus coinciding with observation on the treated shrubs, 
showed natural mortality of less than one percent, and of no significance. 

Limited observations on Ceanothus growing in the wild showed the 
stem-galls to be less abundant than on Ceanothus grown under culti- 
vation. This might indicate that irrigation and other care which these 
ornamentals received under cultivation makes them more attractive 
to this gall insect. 

SUMMARY 

The stem-gall moth, Periploca ceanothiella ( Cosens ) , is a pest of 
ornamental shrubs in the genus Ceanothus. The moths emerge from the 
galls and lay their eggs throughout spring and early summer. There is 
but one generation or annual brood. Three naturally occurring ichneu- 
monid parasites accounted for up to 19% mortality of the maturing 
gall larvae. No hyperparasites were observed. Examination of 40 species 
and varieties of Ceanothus growing in mixed collections at several 
southern California locations showed 2 with heavy occurrence of the 
galls, 4 with moderate occurrence, 13 with light occurrence, and 21 
with no occurrence. This would suggest the planting of the more 
non-susceptible varieties to alleviate or avoid the problem where 
practicable. Dimethoate used as a foliar spray proved highly effective 
against the immature gall larvae, as well as the leaf miner, thrips, and 
mites, but had no apparent effect on the mature gall larvae and pupae, 
or the parasites. Limited observations indicated the galls to be more 
troublesome on Ceanothus growing under cultivation than in the wild. 


REFERENCES CITED 

VAN RENSSELAR, M., and H. E. McMINN. 1942. Ceanothus. Gillick 
Press, Berkeley, California. 308 pp. 

MUESEBECK, C. F. W., K. V. KROMBEIN, and H. K. TOWNES. 1951. 
Hymenoptera North of Mexico Synoptic catalog. Monograph No. 2, U. S. 
Department of Agriculture, Washington, D.C. pp. 186, 188 and 387. 
HODGES, R. W. 1962 . A review of the genus Periploca with descriptions 
of nine new species. Pan-Pacific Entomologist. 38: 83-89. 


Journal of Research on the Lepidoptera 


1(3):19M93, 1963 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
© Copyright 15)63 


LARVAL FOOD-PLANT RECORDS FOR 
SIX WESTERN PAPILIOS 

JOHN F. EMM£L and THOMAS C. EMMEL 

Stanford University, California and Reed College, Portland, Oregon 

Reported on here are some of our recent observations on 
natural and laboratory foodplants used by Papilio eurym-edon Lucas, 
P. indra indra Reakirt, P. indra pergamus Edwards, P. bairdii Edwards, 
P. oregonius Edwards, and P. rudkini Comstock. Some previously re- 
ported foodplants whose authenticity has been doubted are shown to be 
true larval foodplants, while several new plant records for these species 
are reported for northern Oregon and California populations. 

1. Papilio eurymedon. 

After a thorough review of available records, Brower (1958; 1962, 
in litt.) concluded that Pmnus is not a foodplant of this species. The 
present authors (1962), however, recorded this plant as a possible 
foodplant for P. eurymedon, and several observations of eurymedon 
using Prunus seem to be worthy of publishing in view of the previous 
conflicting evidence. 

While collecting in the vicinity of Frazier Park, Kern County, 
California, on June 30, 1962, the senior author observed what appeared 
to be a female Papilio eurymedon fluttering over a small bush of Prunus 
ilicifolia. This observation led to a search of the plant, which yielded one 
second-instar Papilio larva. This larva was brought back to the labora- 
tory and was successfully reared on Prunus lyoni. It pupated on August 
6 and on August 22, it produced a male eurymedon. 

Noel McFarland (1962, in litt.) has also found Prunus ilicifolia, 
as well as Rhamnus crocea, to be foodplants of Papilio eurymedon. At 
Oak Pass in the Santa Monica Mountains of Southern California, he 
states: "Prunus ilicifolia ... is the only plant I have ever found them 
on (beyond second instar) in the wild. I have often collected eggs and 
first instar larvae on Rhamnus croceaP 

At least in California, then, Prunus ilicifolia seems to be a natural 
and fully satisfactory foodplant for Papilio eurymedon. 

2. Papilio indra indra. 

In his Butterflies of North America (1897), Edwards stated that 
Artemisia dracunculoides (Compositae) was a foodplant of P. indra in 
Colorado. Kent Wilson (1961) apparently used this record in the most 
recent publication of foodplants for this species. However, in 1918 J. C, 
Hopfinger- reported that he had never found indra larvae on A. dracun- 
culoides. He did find black Papilio larvae (very probably indra) on an 


191 


192 


EMMEL AND EMMEL 


/. Res. Lepid. 


"umbelliferous species,” on which he also found larvae of P. zelicaon. 
These black "indr a” larvae would not accept A. dracunculoides when 
transferred to it. 

. As we reported in 1962, the foodplant of P. indr a in the Sierra 
Nevada is Pteryxia terebinthina ( formerly Cymopterus terehinthinus ) , 
There are many botanical records of this plant (Dr. Mildred Mathias, 
personal communication) for the area around Brewster, Washington, and 
the "umbelliferous” foodplant found by Hopfinger may well have been 
this species. Don Eff (1962, in litt.) reports the foodplant of indr a in 
the Front Range of Colorado to be Harbouria trachpleura (Umbelli- 
ferae) . 

3. Papilio indr a pergamus. 

The first known foodplant of this subspecies of indra was found by 
Comstock (1928); this was Tauschia parishii (Urn belli ferae) in the 
San Gabriel Mountains. 

Fred Thorne (1962, in litt.) has found pergamus eggs and larvae 
on Tauschia arguta and Lomatium lucidum (Umbelliferae) on Tecate 
Peak, San Diego County, California. 

4. Papilio bairdii. 

Edwards (1893, 1898) found that carrot (Daucus carota) was 
somewhat acceptable to bairdii larvae, while the larvae "thrived” on 
fennel {Foeniculum vulgar e) . However, Brown (1957) states that 
these two plants are unacceptable to bairdii. 

On July 27, 1962, the senior author collected 6 fifth-instar larvae 
and 2 fourth-instar larvae of P. bairdii on Artemisia dracunculoides 
(Compositae) at Barton Flats, San Bernardino County, California. In 
the laboratory, these larvae immediately accepted fennel when placed 
on this plant. Fennel and this Artemisia were eaten with no preference 
for either plant. These larvae pupated, and a male and female adult 
pair emerged on September 9, 1962. 

5. Papilio oregonius. 

On September 1, 1962, both authors collected larvae of P. oregonius 
at Heppner Junction (Gilliam County) along the Columbia River, 
Oregon. These larvae ( 1 second-instar, 2 fourth-instar, and 3 fifth 
instar larvae) were found on Artemisia dracunculoides. In the labora- 
tory, they fed readily on fennel. It was of possible biochemical interest 
to note that the odors of crushed leaves of these plants were similar. 

6. Papilio rudkini. 

The natural foodplant of this species is Thamnosma montana 
(Rutaceae). But this Papilio has also been found on Daucus carota 
(Umbelliferae) in Yuma, Arizona (Bauer, 1955). 

In April of 1962, the senior author collected ten larvae of Papilio 
rudkini on Thamnosma montana in Sentenac Canyon, San Diego 
County, California. These larvae were transferred to fennel in the 
laboratory, which was fairly acceptable to them (20% mortality). All 
refused to eat Citrus, which is very acceptable to Papilio zelicaon. 




PAPILIO FOOD PLANTS 


193 


SUMMARY OF NATURAL AND LABORATORY FOODPLANTS 
RECORDED IN THIS PAPER 


N— natural foodplant; L=:acceptable as laboratory foodplant. 


Papilio eurymedon Rosaceae 

Pmnus ilicifolia (N) 

Prunus lyo-ni (L) 

Rhamnaceae 

Rhamnus crocea (N) 

Papilio indra indra Umbelliferae 

Pteryxia terehinthina (N) 

Harbouria trachpleura (N) 

Previously-recorded Artemisia dracnticnloidcs is dropped in this paper from the group of known 
foodplants of P. indra indra. 


Papilio indra pergamus 


Papilio bairdii 


Papilio oregonius 


Papilio rudkini 


Umbelliferae 

Tauschia parishii (N) 

Tauschia arguta (N) 

Lomatium lucidum (N) 
Compos itae 

Artemisia dracunculoides (N) 
Umbelliferae 

Daucus car Ota (L) 

Foeniculum vulgare (L) 
Compositae 

Artemisia dramunculoides (N) 
Umbelliferae 

Foeniculum vulgare (L) 

Rutaceae 

Thamnosma montana (N) 
Umbelliferae 

Daucus carota (N, buti 
introduced plant) 
Foeniculum vulgare (L) 


LITERATURE CITED 

BAUER, DAVID L., 1955. Notes on the Papilio machaon complex in Arizona. 
Lepid. News 9: 7-10. 

BROWER, LINCOLN P., 1958. Larval foodplant specificity in butterflies 

of the Papilio glaucus group. Lepid. News 12: 103-114. 

BROWN, F. MARTIN, 1957. Colorado Butterflies, p. 215. 

COMSTOCK, JOHN ADAMS, 1928. Records of the Lorquin Entomological 
Society. Bull. Southern Cal. Acad. Sci. 27: 73- 
EDWARDS, WILLIAM H., 1893. Notes on a polymorphic Papilio. Canad. 
Ent. 25: 253-254., 

, 1897j Butterflies of North America, 3 vols. 

- — j 1898. Further observations on Papilio bairdii Edwards. 
Canad. Ent. 30: 11. 

EMMEL, THOMAS C., & JOHN F. EMMEL, 1962. Ecological studies of 
Rhopalocera in a High Sierran community- — Donner Pass, California. 
L Butterfly associations and distributional factors. Journ. Lepid. Soc. 
16: 23-44. 

HOPFINGER, J. C., 1918. Notes on Papilio indra Reakirt. Ent. News 29: 
354-5. 

WILSON, KENT H., 1961. Family Papilionidae. pp. 30-50. In Ehrlich, 
Paul R. and Anne H., 1961. How to Know the Butterflies, Dubuque, 
Iowa: Brown, 262 pp. 


1 (3) :194, 1963 


Journal of Research on the Lepidoptera 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
© Copyright 


COLIAS PHILODICE IN CHIAPAS, MEXICO 

THOMAS C. EMMEL 

Reed College, Portland, Oregon 

COLiAs PHILODICE GoDART has the widest continuous range of any 
Goliad in North America. The species is found from the northern Arctic 
southward to the edge of desert areas in the Owens Valley of eastern 
California, the highlands of northern Arizona, and the Cochise and Rio 
Grande River valleys of New Mexico and Texas. In addition, three 
isolated populations of the species are known in the central mountains 
of Guatemala (Hovanitz, 1950a). The present note extends the known 
distribution of the species to include the republic of Mexico, and gives 
the frequency of white dimorphic females in the single population 
found. 

The Mexican population of C. philodice was discovered in a long 
valley about 12.5 kilometers southeast (on Highway 190) from the 
plaza in San Cristobal de las Casas, central Chiapas. The valley is about 
a kilometer east of the highway at this point. The elevation above sea 
level is 7600-7700 feet, and the rolling mountains are covered with 
pine-oak forest, with grassy areas in the valleys. 

At the time of our visit (March 26-29, 1959), these Colias were 
in full flight, and almost all specimens were fresh. The males were far 
more numerous than the females, but as every female seen was usually 
collected, the data given in Table 1 on relative abundance of the two 
female color phases may be considered fairly accurate considering the 
small number involved (for these dates). The frequency of white 
females in this locality is within the range of frequency (10 to 20%) 
found in the mountains of the western United States (Hovanitz, 
1950b). 

TABLE 1. Frequenqr of color phases of female Colias philodice m central 

Chiapas, Mexico. 

Yellow Females White Females Total Per cent White ± S.£. 

11 2 13 15.38±10.0% 

LITERATURE CITED 

HOVANITZ, W. 1950a. The biology of Colias butterflies. 

I. The distribution of the North American species. Wasmann Biol. 

8: 49-75. 

HOVANITZ, W, 1950b, The biology of Colias butterflies. 

II, Parallel geographical variation of dimorphic color phases in North 

American species. Wasmann J. Biol. 8: 197-219. 


194 


Journal of Research on the Lepiduptera 1 (3) :195-200, 1963 

1140 \V'\ Orange Grove Aie., Arcadia, California, U.S.A. 

© Copyright 1 ^ 6 } 


NOTES ON THE EARLY STAGES OF 
TWO CALIFORNIA GEOMETRIDS 

JOHN ADAMS COMSTOCK 

Del Mar, California 

AETHALOIDA PACKARDARIA 


This moth was described by Hulst in 1888, as Hemerophila 
packardafia. He subsequently created two synonyms for the same insect, 
namely, Selidosema lachrymosa (1898, described from Los Angeles 
County), and Selidosema homopteroides (1901, from Oregon). 

Aethaloida packardaria was pictured by McDunnough in 1920, on 
Plate VII, fig. ii of his Cleorini Studies. 

In 1937 I published a description and illustration of the mature 
larva, from an example taken in Bouquet Canyon, Los Angeles County. 
This year ( 1962) I was able to obtain eggs and rear the species through 
to maturity, which will serve to supplement and amplify the former 
incomplete record. 

Eggs were secured June 27 from captive females. Ceanothus and 
Adenostoma were supplied to them, and oviposition occurred in clusters 
on the tips of twigs. It was noted that the bases of the eggs in each 
cluster pointed outward. 

EGG: 0.8 mm. high, x 0.6 mm. at the widest point. The form is 
subovoid, the base wider and the top tapering to a well rounded and 
smooth surface, with no discernible micropyle. The base tapers inward 
at the circumference and bears a circlet of pearl-like protrusions, about 
20 in number, enclosing a granular depressed floor. 

The surface of the egg above the circlet is finely granular, and does 
not bear ridges. The color, when first laid, is dull olive-green. Later it 
becomes darker. 

Eggs laid June 27 hatched July 2 and 3, 1962. The illustration 
(fig. lA) , shows the egg in lateral aspect. 

FIRST INSTAR LARVA: Length, 2 mm. Width of a typical 
body segment approximately 0.15 mm. 

Head; width, 0.35 mm. Color, jet black except for a narrow trans- 
verse bar on the labrum, which is dull white. The ocelli are black, and 
the setae white. 

The body is predominantly black. The thoracic segments taper from 
the widest next to the head to the third which grades into the 0.15 


195 


196 


COMSTOCK 


/. Res. Lepid. 


mm. wide first abdominal segment. The dorsum is solidly black. Laterally 
on each segment there is a large subtriangulate white spot on each side, 
which extends up over the dorso-lateral edge. 

The legs are black, as are also the prolegs. There are rows of short 
white setae, the most prominent of which is in longitudinal alignment 
above the white spots. 

The first instar larva is shown on figure IB. No attempt has been 
made to accurately map the setae, but preserved larvae are available 
for this purpose. 

SECOND INSTAR LARVA, observed July 7, ’62. Length, 4.5 mm. 
Body width, 0.4 mm., as is also the head width. 

Head, black, as in the first instar. 

The body is a dull maroon over the dorsum. The white triangulate 
spots along the side persist as clumps of round spots in groups of ap- 
proximately three, close together, or fusing with each other. There is a 
transverse line of round white dots on the front margin of the first 
segment. 

The venter is dull maroon. 

This batch of larvae did not thrive well, probably as a result of 
too quick drying of the food plant. Mature larvae were later beaten 
from Adeno stoma and Ceanothus, and were carried through to maturity. 
My previous published description and illustration of the full grown 
larvae (1937) does not need to be repeated, but a drawing is here 
included on figure 1C. 

PUPA: Length, 13.5 mm. Greatest width, 3 mm. Color, black on 
wing cases, thoracic and head portions; red -brown on the abdominal 
segments, with a darker shading on the segmental juncmres, and the 
cauda. 

Two black knobs protrude from the front of the head. The maxillae 
and antennae reach to the wing margins. The cremaster consists of a 
knobbed triangulate black element, from which arises a pair of rela- 
tively long recurved spikes, and along the sides, three short red-brown 
recurved booklets. The spiracles are concolorous with the body, and 
relatively inconspicuous. The pupa is pictured on figure ID. 


BIBLIOGRAPHY 

COMSTOCK, JOHN ADAMS. 1937. Miscellaneous Notes on Western 
Lepidoptera. Aethaloida packmdaria. Bull. So. Calif. Acad. Sci. 36 (3) : 
41-124. PI. 47. 

HULST, G. D. 1888. Hemorophila packardaria. Ent. Amer. Ill: 217. 
HULST, G. D. 1888. Hemerophila packardaria. Ent. Amer. 111:217. 

. 1898. Selidosema lachrymosa. Can. Ent. 30: 194. 

— . 1901. Selidosema homopteroides. Jour. N. Y. Ent. Soc. 

8: 219. 

McDUNNOUGH, J. H. 1920. Studies in North American Cleorini, (Geo- 
metridae) Bull. Dept, of Agric. Canada. 18: 37-38. Plate VII, fig. 11. 


197 


i(}):i9^-200, 196} 


EARLY STAGES 



Fig. 1, Early stages of Aethaloida packardaria (Hulst). A. Egg, lateral 
aspect, enlarged X 50. B. First Instar larva, dorsal aspect, enlarged X 30, 
C. Mature larva, lateral aspect, enlarged X 3. D. Pupa, ventral aspect, 
enlarged X 5. 

Reproduced from water color drawing hy the author. 

PROCHOERODES FORFICARIA 

In 1956 I published notes on the larva and pupa of the geometrid 
moth, Prochoeroides forficaria (Guenee) from larvae obtained in Gila 
County, Arizona. 


198 


COMSTOCK 


/. R<s. Lcpid. 


Dr. McDunnough’s published comments on this species (1940) 
suggest that the Arizona specimens from which I described the larva 
and pupa are closer to form catenulata Grote, and that this "could very 
readily be considered a good species.” 

I have recently reared typical P. forficaria from material taken at 
Del Mar, and the following notes, with illustrations, give a more 
accurate analysis of the species designated 

Prochoerodes forficaria 

Eggs were laid April 10, 1962 in a rearing jar. They were laid on 
their sides, some singly, others in small bunches. 

EGG: When first laid, bright green. Later they became dark, and 
prior to hatching were black. The form was an elongate oval, with about 
16 ridges extending from base to micropyle. There were no horizontal 
ridges or lines. The micropyle was pitted. 

The average length of the egg was 0.9 mm. and the width, 0.5 mm. 

There was considerable variation in size and form, some examples 
being elongate and somewhat less rounded at the ends. 

The eggs hatched April 21 and 22, the larvae usually emerging 
from the side, leaving the remainder of the shell intact. 

Not knowing the food plant, the following were tried; lettuce, 
dandelion, plantain, grass, fuschia, Ceanothus and Cneridium, none of 
which proved acceptable. They finally accepted willow. 

EIRST INSTAR LARVA: Length, 4. mm. Width, 0.2 mm. Head 
width, 0.4 mm. The head is relatively large, and is mottled black and 
brown. Ocelli, black. 

The body is cylindrical, elongate, and very narrow. The ground 
color is slaty gray. There is a broad longitudinal black band on the 
dorsum, with occasional darker spots on some of the segments. The 
lower lateral surface is a translucent gray-straw. The entire ventral 
surface is black. The single pair of prolegs, and anal prolegs, are dark 
on the lateral surfaces and lighter on the ventral surfaces. The setae are 
white, and very minute. 

The first instar larva is illustrated on figure 2B. 

In this phase the larvae were exceedingly active, dropping on a 
thread when touched. 

Larva of 11 mm. in length, — head width, 1.55 mm. 

The body is cylindrical, its entire surface being mottled red-brown, 
on the dorsum there are indefinite quadrate patches of darker brown, 
more clearly defined in the thoracic and caudal areas, but nowhere 
connected sufficiently to form an unbroken band. The true legs are 
yellow-brown, and the prolegs concolorous with the body. The latter 
are prominent and conspicuous. 

MATURE LARVA: Noted on May 5, at the beginning of the 
final instar. Length of larva, 22 mm. Head width, 2.4 mm. 

The body color is light wood brown, the head a shade lighter. The 
body is mottled with darker spots and dashes, and its surface is crossed 
transversely by numerous fine creases and ridges. 



Fig. 2. Egg, larva and pupa of Prochoerodes forficaria. A. Egg, lateral 
aspect, enlarged X 40. B. First instar larva, lateral aspect, enlarged X 24. 
C. Mature larva, enlarged X 5. D. Pupa, ventral aspea, enlarged X 5. 
E. Cremaster, highly magnified. 

Reproduced from water color drawing by the author. 


200 


COMSTOCK 


/. Res. Lepid. 


Middorsally there is an indefinite longitudinal band of brown, 
nowhere clearly defined. Spiracularly there is a pinkish irregular band, 
enlarging around each spiracle, and contracting on the segmental junc- 
tures. Near each spiracle there are two or three round dots. The legs 
are slightly translucent, and the prolegs are concolorous with the body, 
or slightly tinged with yellow. 

There are numerous small papillae scattered over the body, two 
pairs of which in the dorso-caudal area are slightly larger. All papillae 
bear short yellow-brown setae. This is the larval phase that is pictured 
on figure 2C. 

On May 29, the larva measured 30 mm., the head width being 2.4 
mm. as before. The chief difference in this prepupal stage is that there 
is less definite ornamentation, although the pinkish area following the 
spiracles is clearly defined, and the spiracles are more conspicuous, with 
dark rims and a surrounding rim of yellow. 

In the final phase the larva becomes sluggish, and spends much 
time resting on a twig, or the midrib of a leaf, where it is ideally camou- 
flaged. The first example pupated June 9, 1962. 

PUPA: Length, 17 mm. Width through middle portion, 5 mm. 
The color is brown, with the eyes and terminal segments darker. 

The antennae extend slightly beyond the wing margins. The wing 
cases and appendanges are overlaid with fine transverse brown lines 
and dashes. The dorsal surface, and the abdominal segments are heavily 
spotted with dark brown and black. The spiracles are concolorous with 
the body and are inconspicuous. 

The cremaster is formed by a double ovoid extension, narrowing 
down to a clubbed shaft, which ends in two recurved tips that are arched 
latero-ventrally. Near the base of the shaft there are six small yellow- 
brown booklets, three on each side. These arch medially. The pupa is 
illustrated on figure 2D and an enlarged figure of the cremaster on 
figure 2E. 

The first imago emerged June 21, 1962. 

In 1926 Stanley E. Flanders recorded the recovery of three species 
of parasites from larvae of Prochoerodes foriicaria, namely, Amblyteles 
caeruleus (Cr.), A. zebratus (Cr.), and Euplectrus sp. 

We have very little information on the range of P. forficaria but 
Dr, McDunnough’s description of the southern British Columbia form 
combinata suggests that our typical species ranges northward through 
Oregon and perhaps Washington, where it grades into combinata. 

We still need information as to its southern and eastern range. 

BIBLIOGRAPHY 

COMSTOCK, JOHN ADAMS. 1956. Scientific Notes. Prochoerodes for- 
ficaria Gn. Larva, Pupa. Bull. So. Calif. Acad. Sci. 55 ( 3 ) : pp. 180-181. 
FLANDERS, STANLEY E. 1926. Notes on Parasites at Saticoy, California 
During the Year 1925. Pan. Pac. Ent. 2 (7): pp. 157-158. 
McDUNNOUGH, J. H. 1940. New North American Geometridae with 
Notes. 111. Can. Ent. 72(5): pp. 90-103. 


Journal of Research on the Lepidoptera 1 ( 3 ) :201-208, 1963 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 


GEOGRAPHICAL DISTRIBUTION AND VARIATION 
OF THE GENUS ARGYNNIS 
III. ARGYNNIS DIANA 

WILLIAM HOVANITZ 

California Arboretum Foundation, Inc., Arcadia, California, and 
Los Angeles State College, Los Angeles, California 

The Diana fritillary, Argynnis diana Cramer, departs from 
the most usual color pattern for the genus as the pattern is developed 
in most species of the Nearctic and the Palearctic. The male and female 
both have the basal two-thirds of the wings a brownish black. This 
distribution of pigment is in part the reverse of the distribution for 
Argynnis idalia, in which the black pigment is primarily restricted to 
the outer half of the wings. It has already been pointed out (Hovanitz, 
1963) that the distribution of this pigment for idalia is diametrically 
opposed to the ordinary tendencies in other members of the genus in 
which the black pigment is normally distributed in the area of the 
basal regions. This is the pattern arrangement for Argynnis diana, A. 
cyhele, A. leto, A. nokomis and others in North America which show an 
extension of basal melanism. The remainder of the wings shows the 
usual pattern arrangement of black dots and black striping which is 
typical of the genus Argynnis in any part of the world. In the male, 
however, these black markings are greatly diminshed and restricted in 
most cases, especially on the hind wing, so that the outer one-third of 
the wings on the upper side are nearly a solid brownish-orange color. 
On the female this restriction of the black does not occur. It is in fact 
greatly extended so that not only is the outer margin all black, but the 
black-veined stripes are wide and extend from the basal black to the 
margin, and the black dots are large and converge to form black stripes 
running nearly from the anal margin to the inner margin at least on 
the fore wing. The part of the wings then remaining that would nor- 
mally be a brown color in Argynnis is an irridescent blue. On the face 
of it, the female diana is a black butterfly with marginal and sub- 
marginal rows of blue spots. This dimorphism of the sexes of A. diana 
is the greatest expression in North American Argynnis of this type of 
variation. It is, however, nearly duplicated by some races of Argynnis 
nokomis, to which it may be closely related (fig. 1 ) . 

The under side of the wings also shows a very different aspect than 
the usual Argynnis pattern. This alteration of the pattern is more than 
the usual general suffusion of color over the wing but effects the pattern 


201 


202 


HOVANITZ 


/. Res. Lcpid. 



i^ssi 

in»!! 

**#£::****»• 


!!{******••••< 

!!!!!•••••••••• 

lISSS!!*****«*»‘t ^ 


p 

niK!!****«> 

£/■?*•«»«».» 

f 


3(ppii: 

LvErs:?""' 

iJHBBlS!!****' 

HHsili-f 

i^Hl!*****- 

rai::*!” 


Fig. L Argynnis diana, upper side. Top, male; bottom, female. The 
blue is irridescent and may show only in certain directions of certain quality 
of light. ( See fig. 2 ) . 


I (}): 20 I - 2 o 8 , 1963 


ARGYNNIS DISTRIBUTION 


203 



Fig. 2. Same as fig. 1, only under side. $ Salem, Roanoke Co., Vir- 
ginia. June 13, 1937. C. W. Gottschalk. 2 Montgomery Co., Virginia. Aug. 
16, 1902. E. A, Smyth, Jr. 


204 


HOVANITZ 


/. Res. Lepid. 


elements in such a way as to be most certainly of fundamental embryo- 
logical nature. By this is meant the almost complete obliteration of the 
pattern elements on the underside of the hindwings in both the male 
and the female, with a slight degree of the same on the fore wings. In 
this development, A. diana is also most directly opposed to A. idalia in 
which the spots and pattern elements are well developed, even more so 
than in many other Argynnis. The pattern observed on the diana hind 
wing underside looks as though all markings had been wiped clean 
between the most marginal row of small spots and the usual Argynnis 
row of large silvery spots, substituting therefore, a uniform bluish 
black on the female, light brown in the male; likewise for the area 
from the base of the wing outward to the same row of spots nearly all 
markings of the usual Argynnis type appear to have been wiped clean 
except for a few minor points and substituting therefore a brownish- 
black on the female and light brown in the male. A small part of the 
central row of silvery spots of the usual Argynnis pattern remains, along 
with a small part of the cell spot and most marginal row in both the 
male and the female. The inner two-thirds of the fore wings on the 
under side in both sexes are similar to the usual Argynnis pattern. 
Interestingly enough, there is blue irridescence in this part of the wings 
of the male, even though the upper side of the male wings do not 
show it. ( fig. 2 ) 

A similar development of the pattern elements on the under side of 
Argynnis wings occurs in the groups related to A. paphia, A. pandora, 
A. laodice, A. childreni, etc. of Eurasia. These are the groups which 
have a distributional range through areas of higher rainfall, and thus 
more humid air, combined with higher summer temperatures. Some 
membrs of these groups also have a development of sexual dimorphism 
similar to that of A. diana and A. nokomis, even to the extent of having 
blue or bluish females while the male is the usual orange-brown color. 
More complete correlations between wing color, pattern type and insect 
size with climatic differences in various parts of the distributional range 
of Argynnis will come in a later issue. 

The geographical distribution of Argynnis diana is a rather re- 
stricted one (fig. 3), hardly being more than 10° from the most 
northern part of its range to the most southern part and 15° from the 
most easterly to the most westerly part of its range. This area in miles 
is abou 1000 miles from east to west and three to four hundred miles 
from north to south. Actually, the present known distributional range 
is shaped somewhat like a triangle, with an acute angle in the far west 
in Arkansas. Except for a collection in western Pennsylvania which 
may be inaccurate, the species is known only as far north as southern 
West Virginia and central Virginia. The most southern localities are 
southern South Carolina and northern Georgia and an area of north- 
central Arkansas. There are records of the species from southern 
Illinois. It is likely that they may be found in southern Missouri, southern 


(});20i-2o8, 1^6} 


ARGYNNIS DISTRIBUTION 


205 


NORTH AMERICA 



Fig. 3. Map showing the North American distribution of Argynnis 
diana and A. nokomis. Note the wide expanse of isolation between these two 
closely related butterflies; also the wide isolation between parts of the range 
of A. nokomis. Details on the geographical variation of nokomis will come in 
the next number in this series. 


206 


HOVANITZ 


/. Res. Lepid. 


Indiana and southern Ohio in addition, though I have not seen specimens 
from these locations. 

This species, despite its striking colors and size, is a relatively rare 
species due to the limited areas in which it lives. There is no evidence 
that at any time in the past the species was any less local than it is now, 
even though it is said that the deforestation of many parts of the east 
have contributed to its demise in many areas (Clark, 1951). The 
species exists in parts of North America away from the most heavily 
inhabited regions and therefore collectors have not had as much experi- 
ence with it as with other species. Undoubtedly, the restricted habitat 
of the larvae contributes most to its own restricted range and the factors 
that are essential for larval environment are not known. Judging how- 
ever from the habitats preferred by the adults and especially the females 
which is usually highly indicative, the species finds its home in areas 
of cold seepages and streams on the coastal plain, or in deep, damp, 
well wooded valleys or ravines, or on damp wooded mountain sides 
where there are cold streams (Clark, 1951). This corresponds to the 
habitat of A. nokomis, the western related species (fig. 1), which 
occurs in dry desert regions but only along cold streams where there 
is an abundance of boggy vegetation and locally a "wet-tropical” micro- 
habitat as at Round Valley, Inyo County, California. This type of 
habitat is very local, not only in the desert regions of the west but also 
apparently in the east and is probably more responsible for the restric- 
tion of range of diana than the removal of the forest cover over exten- 
sive areas of the eastern parts of North America. 

Clark (1951) gives the most complete description of the distribu- 
tion, variation and habits of this species than has ever been given. The 
following is from his description : 

"The first males to appear are smaller than those that emerge later, 
and the males from Highland County and from the higher latitudes 
in the mountains farther south are always small, resembling the earliest 
males from other regions. 

"On the Coastal plain the males occasionally have on the under side 
of the hind wings at the end of the cell a conspicuous silver spot 
bordered inwardly and outwardly by black lines, corresponding to the 
silver spot in the same position in S. cybele though smaller, seldom 
reaching more than halfway across the cell, and the silver markings on 
the outer portion of the hind wings may be enlarged. Occasionally on 
the upper surface of the fore wings there are broad light dashes in the 
black ground color beyond the cell in the interspaces between veins 3 
and 4, 4 and 5, 5 and 6, corresponding to the light dashes on the under 
side though with indefinite borders. So far as we have seen, these 
features are confined to males from the Coastal Virginia Plain. The 
majority of the males from the Coastal Plain, however, do not differ 
in any way from others from the mountains. 

"Dr. Henry Skinner has pointed out that the males differ materially 


l(})t20l-20i, 1^6} 


ARGYNNIS DISTRIBUTION 


207 


in the number and size of the black spots on the upper surface of the 
hind wings. We have seen males, both from the Coastal Plain and the 
mountains, with the light border of the hind wings almost immaculate. 
There is some variations in the size of the black spots on the fore wings, 
and veins 2, 3, and 4, may be narrowly or rather broadly infuscated. 
The contrast between the dark basal and light outer portions of the 
under surface of the hind wings is sometimes accentuated. These varia- 
tions seem to bear no relation to locality, 

"Dr. Skinner said that females from eastern Tennessee, western 
North Carolina, and southern Illinois are larger than those found in 
Virginia; but the females from the lower altitudes in southwestern 
Virginia appear to be quite as large as any from farther south. Dr. 
Skinner pointed out that the females vary a great deal in the degree of 
silver beneath and also in the band of large bluish or greenish spots 
on the hind wings above. In sorhe specimens these spots are large, and 
in others they are confined to a small area around the black spots. 
The number and size of the cream-colored or white spots on the upper 
side of the fore wings is also quite variable.” 

It appears from my own observations as well as from those of others 
that there is quite a lot of variability in this species; most of these 
variations however appear to be individual rather than population in 
scope, with the exception of those that are directly related to the eleva- 
tion differences described by Clark. Occasionally the blue irridescence 
has been changed to green. Statistical study of large samples seems to be 
called for to bring this variation out of the realm of guesswork. 

The species flies over a long season, males coming out in the middle 
of June and females slightly later with a height of emergence in the 
middle of August and extending into September. 

Most authors in discussing this species in handbooks comment on 
mimicry between this species and some other species which inhabit the 
same general geographical area. Ehrlich (1961) states "The female 
presumably mimics the distasteful Aristolochia swallowtail, Battus 
philenor,” Klots (1951) states "This is cited as a case of mimicry, the 
supposed model being the black and blue Papilio philenor, a swallowtail 
supposedly distasteful to birds.” Scudder (1889) considers the female 
is a case of parastatic mimicry with Basilarchia aslyanax. It is fortunate 
that most authors leave the case in doubt as most all situations of this 
sort are wild grasps at straws in the wind. The relationships in appear- 
ance of many butterflies in the same general geographical area, such as 
these three, A, diana in the east and A. nokomis in the southwest with 
Papilio philenof and Basilarchia astyanax are exceedingly interesting 
and deserve some intensive study. Too often these studies are superficial 
and involve some transference of a man’s way of thinking to a butterfly 
or a bird. As purely a suggestion, it appears that a more satisfactory 
solution may be found in the physiological relationship of the butterfly 
with its physical environment, in which the colors are related in some 


208 


HOVANITZ 


/. Res. Lepid. 


way to light reflection from the butterfly’s wings (U.V. light perhaps) 
which reflections are developed in the butterflies living in a warmer 
or more humid environment. 


LITERATURE CITED 

CLARK, A. H. 1951. The Butterflies of Virginia. Smithsonian Misc. 
Collections, 116(7): 1-95. 

EHRLICH, PAUL and ANNE H. 1961. How to know the butterflies. 
Brown, Dubuque, Iowa. 

HOVANITZ, WILLIAM. 1963. Geographical distribution and variation 
of the genus Argynnis. J. Res. Lepid. 1(2): 117-123. 

KLOTS, A. H. 195 L Field guide to the butterflies. Houghton Mifflin, 
Boston. 

SCUDDER, S. 1889. The Butterflies of the Eastern United States. Cam- 
bridge, Mass. 


Journal of Research on the Lepidoptem 1 (3 ) :209-221, 1963 

1140 W, Orange Grove Ave., Arcadia, California, U.S,A. 

© Copyright 196 } 

THE GENERIC, SPECIFIC AND LOWER CATEGORY 
NAMES OF THE NEARCTIC BUTTERFLIES 
PART 2 — The Genus CoUas 

PADDY. McHENRY 

1 032 E. Santa Amta^ Burbank, California 

The names included io this genus are presented in the same 
manner as were those of the genus Pieris in the first part of this series 
{]om. Res. Lepid. 1(1) :63-71, 1962). 

W, Hovanitz has separated the members of the genus into specific 
groups as best he felt was possible in ¥iew of the extensive hybridiza- 
tion in the genus. ‘ 

Tlie genus Zerem Hiibner, although considered a subgenus of 
Colias by some students, will be dealt with separately in a subsequent 
part. 

The data for chfysojheme Esper has been listed under the eurytheme 
group purely as a convenient place in which to give this reference; 
should a relationship be established between chrysotheme and eurytheme, 
the former must, of course, be accorded priority. 

The name notatus has been presented in the philodice group under 
the authorship of Megerk and also under the authorship of Clark and 
Clark because of its present uncertain nomenclatorial status. 

LIST OF GENERIC NAMES USED OR AVAILABLE FOR 

COLIAS, 

COLIAS Fabricus. 

Type: hyde Linnaeus. 

EURYMUS Horsfield. 

Type: h'yde Linnaeus. 

COLIAS FABRICIUS. 1807 [before 19 Deep Io K, Illiger. Magazin fur- 
Insekteokunde 6: 284, oo. 24, Includes among others ^Pap . . . 
Hyde’* and Rhamni" both considered to be of Linnaeus, 1758. 
Type. P[apiUo\ D^anms]. {Candidas'] hyde Linnaeus. 1758. Syst. Nat. 
10th. Ed. 1: 469, no. 71. 

Type Selection. 30 Sept. 1943. Opinions & Declarations Rendered by 
the International Commission on Zoological Nomenclature 2(13): 
109-121, The type is declared to be hyde Linnaeus, 1758. This 
declaration invalidates the selection of ^Colias Rhamni, Fab,” (con- 
sidered to be rhamni Linnaeus, 1758) by Latreille, 1810, Consid. 
Gen. Anim. Crust. Arach. Ins, : 440. 

EURYMUS HORSFIELD, (Swainson Ms. name). 1829. [24 June]^. 

Catal. Lepid. Ins. Contained Mus. Honor. East Indies Company 
(2) : 134. Gives only ^^Colias Hyale” which is considered to be 
hyale Linnaeus, 1758. The name is a homonym of Eurymas Rafin- 
esque, 1815^. 

Type. P[apilm]. D\anaus]. [Candidm] hyale Linnaeus. 1758. Syst. Nat. 
10th. Ed. 1 : 469, oo. 71. 

Type Selection. Horsfield. 1829 [24 June]^. Ibid. (2) : 134. "Mr. Swain- 
son gives Colias Hyale as the type of this subgenus . . 


209 


210 


McHENRY 


/. Res. Lepid. 


LIST OF SPECIES AND LOWER CATEGORY NAMES USED OR 
AVAILABLE FOR COLIAS 


1. COLIAS BEHRII W. H. 
EDWARDS. 

behrii W. H. Edwards. 
ccmescens (J. A. Comstock). 

2. COLIAS BOOTHII CURTIS. 
boothii Curtis. 

chione Curtis. 

3. COLIAS EURYTHEME 
BOISDUVAL. 

alba Strecker. 
alba Strecker. 
amphidusa Boisduval. 
ariadne W. H. Edwards. 
californiana Menetries 
chrysotheme (Esper). 
eurytheme Boisduval. 
flam Strecker. 
fumosa Strecker. 
intermedia Cockerell. 
keewaydin W. H. Edwards. 
pallida Cockerell. 
pallida Cockerell. 
pallida Cockerell. 
rudkini (Gunder). 
unicitrina (Gunder). 

4. COLIAS GIGANTEA 
STRECKER. 
gigantea Strecker. 
harroweri Klots. 

marjorie Chermock & Chermock. 
ma^i Chermock & Chermock. 
pelidneides Staudinger. 

5. COLIAS HECLA LEFEBVRE, 
chrysothemoides Verity. 
glacialis M’Lachlan. 

hecla Lefebvre. 
hela Strecker. 
palamedes Hemming. 
pallida Skinner. 

6. COLIAS MEADII W. H. 
EDWARDS. 

elis Strecker. 

meadii W. H. Edwards. 

medi (Gunder). 

7. COLIAS NASTES 
BOISDUVAL. 
alias ka Bang-Haas. 
cocandicides Verity. 
gueneei Avinoff., 
harperi (Gunder). 
moina Strecker. 
nastes Boisduval. 
ob scut at a Verity. 
palliflava (McDunnough) . 
rossii Guenee. 


streckeri Grum-Grshimailo. 

subarctica (McDunnough) . 
thula Hovanitz. 

8. COLIAS OCCIDENTALIS 
SCUDDER. 

alba Strecker. 
alberta Bowman. 
alexandra W. H. Edwards. 
astraea W. H. Edwards. 
bar bar a Henry Edwards. 
Christina W. H. Edwards. 
chrysomelas Henry Edwards. 
edtvardsii W. H. Edwards. 
emilia W. H. Edwards. 
harfordii Henry Edwards. 
hatui (Barnes & Benjamin). 
krauthii Klots. 
lambillioni Dufrane. 
martini (Gunder). 
occidentalis Scudder. 
pallida Cockerell. 
pallida Cockerell. 
pallidissima Bowman, 
shastae (Barnes & Benjamin). 
weaverae ( Gunder ) . 

9. COLIAS PALAENO 
(LINNAEUS). 
chippewa W. H. Edwards. 
helena W. H. Edwards. 
kohlsaati (Gunder). 
lapponica Staudinger. 
palaeno (Linnaeus). 
philomene (Hubner). 
werdandi Herrich-Schaffer. 

10. COLIAS PELIDNE 
BOISDUVAL & LECONTE. 
intertor Scudder. 

isni (Barnes & Benjamin). 
labradorensis Scudder. 
laurentina (Scudder). 
minisni Bean. 
mira Verity. 

moeschleri Grum-Grshimailo. 
nepi (Barnes & Benjamin). 
neri (Barnes & Benjamin). 
pelidne Boisduval & LeConte. 
skinneri Barnes. 
solivaga W. H. Edwards. 

1 1 . COLIAS PHILODICE 
GODART. 

alba Strecker. 
alba F. Chermock. 
albida (F. Chermock), 
albinic W. H. Edwards. 
ant hy ale (Hubner). 


(});209~22l, 196} 


NAMES — Pt. 2 


211 


autumndis Cockerell. 
ehrmanni F. Chermock. 
eriphyle W. H. Edwards. 
europome ( Haworth ) . 
hagenii W. H. Edwards. 
hybrida Strecker. 
inversata Nakahara. 
kootenai Cockle. 
laurae (F. Chermock). 
lutetincta Wolcott. 
melanic W. H. Edwards. 
melanic W. H. Edwards, 
minor F. Chermock. 
miscidice (Scudder). 
nig Strecker. 

nigricosta (F. Chermock). 
nigfidice ( Scudder ) . 
nigrina Strecker, 


nigrofasciata Reiff. 
notatus (Mergerle). 
notatus (Clark & Clark). 
pdlidice ( Scudder) . 
philodice Godart. 
plicaduta Nakahara. 
raritus (Gunder). 
reducta Dufrane. 
rothkei Reiff. 
s errata (F. Chermock). 
suffusa Cockerell. 
virida Strecker. 
vitahunda Hovanitz^ 


12. COLIAS SCUDDERII 
REAKIRT. 
flavotincta Cockerell. 
ruckesi Klots. 
scudderii Reakirt. 


1. COLIAS BEHRII W. H. EDWARDS. 

hehrii, Colias W. H. Edwards. Oct. 1866. Proc. Ent. Soc. Phila. 6 
(pp. 201-208): 201. "From 2 $, I $, received from Dr. Behr, 
and taken among the Yo Semite mountains at an elevation of about 
10,000 feet above the sea.” No dates given. Spelled behri by J. A. 
Comstock, 20 May 1925, Bull. Sou. Calif. Acad. Sci. 24(1): 3. 
canescens, Eurymus behri J. A. Comstock. 20 May 1925. Bull. Sou. 
Calif. Acad, Sci. 24(1) : 3. "Type [$], Tioga Pass, Yosemite, Calif., 
Aug. 4 , 1922 . Six paratypes [$ $]. Same place and date. In the 
collection of the Southwest Museum.” 

2. COLIAS BOOTHII CURTIS. 

chione, Colias Curtis. 1835. In J. Ross. Append. Narr. 2nd Voy. 

Search North-West Pass. (Nat, Hist. Sect.), pp. Ixvi-lxvii, no, 11; 
plate A, fig. 6. d & $ described. ”... they were captured from 

the I4th of July to the 13th of August, 1830, and on the 19th 

were in a very wasted state; on the l4th of July, of the following 
year, one Colias only was taken.” No locality nor series data given 
here. 

boothii, Colias Curtis, 1835. In J. Ross. Append. Narr. 2nd. Voy. 

Search North-West Pass. (Nat. Hist. Sect.), pp. Ixv-lxvi, no. 10; 
plate A, figs. 3-5. d & $ described. No locality, series data nor 
dates given here. Spelled boothi by Verity, 1905-1911, Rhopal. 
Palaearaica, p. 238. 

3. COLIAS EURYTHEME BOISDUVAL. 

alba, Colias chrysotheme eurytheme Strecker. 1878 [Sept. - Ort.]"^. Butt. 
& Moths N. Amer., Compl. Syn. Catal., p. 83, no. 60b. Cites $ 
figures in W. H. Edwards, Butt. N. Amer. [I]: plate Colias III, figs. 
5-6, 1869. No locality, series data nor dates given here. 
alba, Colias chrysotheme keewaydin Strecker. 1878 [Sept. - Oct.]"^. Butt. 
& Moths N. Amer., Compl. Syn. Catal., p. 83, no. 60e. Cites $ 
figures in W. H. Edwards. Butt. N. Amer. [1]: plate Colias IV, figs. 
8-9, 1869. No locality, series data nor dates given here. 
amphidusa, Colias Boisduval. 1852 [Aug.]^ Ann. Soc. Ent. France. 2nd. 
Ser. 10(2): 286-287, no. 14. d & 9 described. "Du nord de la 
Californie.” No series data nor dates given. Spelled amphidona 
by W. H. Edwards, 1869, Butt. N. Amer. [1]: [45]. 
ariadne, Colias W. H. Edwards. Jan. 1870. Trans. Amer. Ent. Soc. 3(?) 
sign. 2: 12-13. ”1 d, 1 9 , from the collection of Dr. Behr, and 
taken at Mokeluma Hills, California.” No dates given. 


212 


McHENRY 


/. Res. Lepid. 


Californian, Colias edusa Menetries. Dec. 1855. Enum. Corp. Anim. 
Mus. Imper. Acad. Sci. Petropolitaoae. Class. Ins. Ordo Lepid. ( 1 ) : 
80, under no. 253. "Ces caraaeres indiques ci dessus se reproduisent 
constamment sur environs 6 males et 4 femelles, tous rapportes de 
la Nouvelle California par notre zele voyageur Wosnesensky.” No 
dates given. 

chrysotheme, Dan[aus]. Can{didus\ Esper. [1781]*. Die Schmetterlinge 
1(2) heft 3: 89-90, no. 131; plate 65, figs. 3-4. d & $ described. 
No locality, series data nor dates given. Note: This name is included 
here only for convenience. 

eurytheme, Colias Boisduval 1852 [Aug.]*. Ann. Soc. Ent. France. 2nd. 
Ser. 10(2): 286, no. 13. "Commune dans toute la Californie. 
Elle habite aussi le Mexique et quelques parties des Etats-Unis.” 
No sex, series data, nor dates given. Spelled enegthenu by W. H. 
Edwards, 1872, in F. V. Hayden, Prel. Kept. U. S. Geol. Surv. Mont. 
& Port. Adj. Terr. 5th. Kept. Progress, p. 466. 
flava, Colias chrysotheme eurytheme Strecker. 1878 [Sept. - Oct.]"^. Butt. 
& Moths. N. Amer., Compl. Syn. Catal., p. 83, no. 60c. 2 described. 
"Mus. Strecker." No locality, series data nor dates given. 
fumosa, Colias eurytheme Strecker. 1900 [Mar. 9th-13th]^. Lepid., 
Rhopal. & Heter., Indig. & Exot. Suppl. No, 3: 19. "One $, 
Colorado . . No date given. 

intermedia, Colias eurytheme Cockerell. Mar. 1888 [after 9th]®. West 
Amer. Scientist 4(35): 41-43 (in pt.). 2 described. "In this 

locality (Swift Creek, Custer Co., Colo[rado] , . No series data 
nor dates given. 

keewaydin, Colias W. H. Edwards. [Sept. 1869, on or before 13th]'^, 
Butt. N. Amer. [I](4) : [47]-[49]; plate (15), figs. 1-9. d & 2 
described. '^Found in the valley of the Mississippi from Nebraska 
and Illionis to Texas and westward to the Pacific, occupying much 
the same region as Eurytheme, but apparently less common and 
more local than that species. Also occasionally found in the Middle 
States and Canada." "There appear to be two broods during the 
year, the insects being most abundant early in the spring, in faa, 
before Eurytheme makes its appearance.” No series data nor dates 
given. 

pallida, Colias eurytheme Cockerell. Nov. 1887. West. Amer. Scientist 
3(31): 217. 2 described. "... near West Cliff, Custer Co., 

Colorado.” No dates given. 

pallida, Colias eurytheme intermedia Cockerell. Mar. 1888 [after 9th]®. 
West. Amer. Scientist 4(35): 41-43 (in pt.), "... Whitish 
females occur of . . . intermedia, these may be called pallida.” No 
locality, series data nor dates given, 

pallida, Colias eurytheme keewaydin Cockerell. Mar. 1888 [after 9th]®. 
West. Amer. Scientist 4(35): 41-43 (in pt.). . . Whitish 

females occur of . . . keewaydin . . . these may be called pallida.” 
No locality, series data nor dates given. 
fudkini, Eurymus eurytheme Gunder. 31 Dec. 1932. Can. Ent. 64(12) : 
211 -21 k. "Holotype - 2 . . . San Marino, Los Angeles Co., Calif., 
May 23, 1931. Type in Author’s coil.” 
unicitrina, Eurymus eurytheme amphidusa Gunder. 8 May 1924. Ent. 
News 35(5): 158; plate 2, fig. J. "Holotype (Author’s Coll.) 
Upland, Los Angeles County, California; August 2, 1921.” 

4. COLIAS GIGANTEA STRECKER. 

gigantea, Colias pelidne Strecker. 1900 [Mar. 9th"13th]^. Lepid., 
Rhopal. & Heter., Indig. & Exot. Suppl. No. 3: 19. "Five $ S, 
1 2 2, west coast Hudson Bay, above Fort York, Archdeacon 

Kirby.” No dates given. 


t(}):ao$-22ig 1^6} 


NAMES ™ Pt. 2 


213 


hatfoweri, Colias gigantea Klots. 22 Mar. 1940. Amer. Mus. Nov. No. 
1054: 4-6. "Type Lot All are from Clear Creek, vicinity Lower 
Green River Lake, Sublette Co., Wyo[ming]., alt. 8400 ft. Holotype 
male, three male and two female paratypes, August 3, 1935 • . 
allotype female, nine male and one female paratypes, July 16, 
1939...” 

marjofie, Colias Christina mayi Chermock & Chermock. 30 Apr. 1940. 
Can. Ent. 72(4): 82. "Holotype - $, June 26, 1933, Riding 
Mountains, Manitoba. Paratypes - 1-50, same locality.” Page 8N 
"... holotypes will be deposited in Canadian National Collec- 
tion . . .” 

mayh Colias Christina Chermock & Chermock. 30 Apr. 1940. Can. 
Ent. 72(4): 81. "Holotype - d, July 1, 1933, Riding Mountains, 
Manitoba. Allotype - 2, July 1, 1933, same locality. Paratypes - 
1 to 150, same locality.” Page 81: " . . . holotypes will be 
deposited in the Canadian National Colleaion and the allotypes 
in the Carnegie Museum.” 

pelidneides, Colias palaeno Staudinger. May 1901. In Staudinger & 
Rebel. Catal. Lepid. Palaearct. Faun. 3rd. Ed., p. 15, no. 86i. $ 
described. "Amer. bor. (Hudson Bay, Alaska).” No series data 
nor dates given. 

5. COLIAS HECLA LEFEBVRE. 

chrysothemoides, Colias hecla Verity. 1905-1911. Rhopal. Palaearctica: 
p. 356; p. xxxvii; plate LXXI, figs. 22-23; plate LXXI explanation 
page, figs. 22-23. "22 . . .d (Territoire de Barren, 114°, 30’ long. 
O., 67° 40’ lat. N., Am. boreale).” "23 . . .9 ([same as 22]).” 
Collection of British Museum. No series data nor dates given. 

glacialis, Colias hecla M’Lachlan. 23 May 1878. Jour. Linnean Soc. 
(London), Zoology 14(74): 108. "Two $ and one $ from lat. 
81° 45’, August 12th, 1876, and one 2 from Hales Sound, lat. 
79° . . . also a much crippled $ . . . from Discovery Bay, July 
18th, 1876 . . .” 

hecUf Colias Lefebvre. 1836 [after 6 Apr.]’°. Ann. Soc. Ent. France 
5(?)': 383-387; plate 9, figs. 3-6. ^ & 2 described. " V 
Islande est la patrie . . .”. No series data nor dates given. 

hela, Colias Strecker. Sept. 1880. Bull. Brooklyn Ent. Soc. 3(5) : 33-34. 
"... the present descriptions are from 1 d and 2 2 2 ; a few more 
were taken, but these are all I have had the opportunity of exam- 
ining. They were captured ... a considerable distance above Fort 
Churchill on west coast of Hudson’s Bay; . . .”. No dates given. 

palamedes, Colias hecla Hemming. May 1934. Stylops, Jour. axon. 
Ent. (5) : 98, no. 17. A new name for Colias hecla pallida 
Skinner which he states is a homonym of Colias erato pallida 
Staudinger (1861, in Staud. & Wocke, Cat. Lepid. Europ., p. 3). 

pallida, Colias hecla Skinner (Skinner & Mengel Ms. name). 1 Mar. 
1892. Ent. News 3(3) : 49; plate 2, fig. 4. 2 figured. "... from 
Greenland.” No date given, A homonym of Colias erato pallida 
Staudinger (1861, in Staud. & Wocke, Cat. Lepid. Europ., p. 3). 
Spelled palida by Holland, 1931, Butt. Book, rev, ed., p. 298. 

6. COLIAS MEADII W. H. EDWARDS. 

elis, Colias Strecker. 10 Mar. 1885. Proc. Acad. Nat. Sci. Phila. [37] 
(sign. 2) : 24. Description concluded on page 25 (sign. 3) : 7 Apr. 
1885. "Taken ... at an elevation of 10,000 feet, on the summit 
of "Kicking Horse Pass,” in the Rocky Mountains, between Alberta 
Territory and British Columbia, at the boundary between the 
United States and the British possessions, about 300 miles north 
of Montana.” "Capt. Geddes took about fifteen examples, all 
females ...” No dates given. 


214 


McHENRY 


/. Res. Lcpid. 


meadii, Colias W. H. Edwards. Feb. 1871. Trans. Amer. Ent. Soc. 
3(.^) sign. 34: 267-268. 6 & $ described. No locality, series 

data nor dates given. In his Butt. N. Amer. [1]: 59, 1868-1872, 
he says: "Taken in Colorado ... in the month of July, 1871.” He 
speaks of Mr. T. L. Mead taking 12 specimens on the "divide,” 34 
specimens on "Mt. Lincoln” and 1 specimen at "Kenosha House.” 
Spelled meadi by Holland, 1931, Butt. Book, rev. ed., p. 297. 

medi, Eurymus meadii Gunder. 30 June 1934. Can. Ent. 66(6): 125. 
"Holotype - $, . . . Beckenridge Peak (11,000 ft.), Empire County, 
Colorado, Aug. 8, 1919. Type in Author’s coll.” 

7. COLIAS NASTES BOISDUVAL. 

aliaska, Colias nasPes Bang-Haas. 15 Aug. 1927. Horae Macrolepid. 
Reg. Palaearctica 1(3) : 41. Figs. 24-25 on plate 5 were published 
later. "Habitat: Alaska: Ramport 8. Juli, ...,3d 2 $ Spelled 

alaskae by McDunnough, 1938, Mem. Sou. Calif. Acad. Sci. 1: 8, 
no. 52e. 

cocandicides, Colias nastes rossii Verity. 1905-1911. Rhopal. Palaearctica; 
p. 355 (as rossii); p. xxxvii (as cocandicides); plate LXXI, figs. 7-9; 
plate LXXI explanation page, figs. 7-9 (as rossii). "7 . . . d 
(Territoire de Barren, 114°, 30’ long. 0., 67° 40’ lat. N., Am. 
Iwreale)”. "8 . . . $ ([same as 7])”. "9 . . . d ... ([same as 7 
& 8]) ”. No series data nor dates given. 

gueneei, Colias nastes Avinoff. 10 Dec. 1935. In Holland & Avinoff. 
Mem. Carnegie Mus. 12(2): sect. 5, sub-sect. 2: 13-14, no. 10; 
& page 32; plate 27, figs. 1, 11, 21-22, 25-27, 31-32. d & $ 
described. Locality: Southampton Island. No series data nor dates 
given. 

harperi, Eurymus nastes moina Gunder, 31 Dec. 1932. Can. Ent. 64(12) : 
278. "Holotype - 2 , . . .; Ft. Churchill, Manitoba, Can[ada]., July 
19, 1932. Type in Author’s coll.” 

moina, Colias Strecker. Sept. 1880. Bull. Brooklyn Ent. Soc. 3(5): 34. 
d & 2 described. "Described from a large number of examples 
mostly males.” ["a considerable distance above Fort Churchill on 
West Coast of Hudson’s Bay” . . .] (taken from Strecker’s text 
of hela, p. 34) . 

nastes, Colias Boisduval. [1832]”. Icon. Hist. Lepid. Nouv. Peu Connu 
(?): plate 8, figs. 4-5 (d). Pages 245-246 published later 
( d & 2 ) . "Elle se trouve au Cap-Nord et en Islande. M. Sommer 
m’en a aussi envoye plusieurs individus pris au Labrador.” No series 
data nor dates given. 

obscurata, Colias nastes streckeri Yenty. 1905-1911. Rhopal. Palaearctica; 
pp. 354-355 (in pt.); p. xxxvi; plate LXXI, fig. 6; plate LXXI 
explanation page, fig. 6. "6 . . . d ... (Lake Louise, Alberta)”. 
No date given, 

palliflava, Eurymus nastes streckeri McDunnough. 25 July 1927. Can. 
Ent. 59(7): 153-154. d described. "... higher slopes of Mt. 
McLean (above 6500 ft.) on July 12 . . .”. ". . . type No. 2425 
in the Canadian National Collection, Ottawa”. 

rossii, Colias nastes Guenee. 12 Oa. 1864. Ann. Soc. Ent. France. 
4th. Ser. 4(2) : 199-200. d & 2 described. "... trois individus , .”. 
No locality nor dates given here. Spelled rossi by Holland, 1931, 
Butt. Book, rev. ed., p. 299. 

streckeri, Colias nastes Grum-Grshimailo. 30 Apr. 1895. Horae Soc. 
Ent. Rossicae 29 (1-2) : 290-291, no. 2. d & 2 described. "Specimen 
unum hujus varietatis sub nomine ^'Colias Behrii?” a lepidopterologo 
germanico D-re 0. Staudinger anno 1891, quattuor specimina, in 
provincia Alberta ad Laggan collecta, a lepidopterologo americano 
Dom. H. Strecker, cujus in honorem hanc forman nominavi, accepi”. 


I (}) ‘.20^-221, 1^6} 


NAMES --- Pt. 2 


215 


subarctica, Eurymus nastes McDunnough. 1 Dec. 1928. Cao. Ent. 
60(11): 270^271. "Holotype. ^ Bernard Harbour, N. W. T., 
Aug. 9, 1915, . . . , No. 2863 in the National Collection, Ottawa. 
Allotype. - 2 , same data. Paratypes. - 4 d , 4 $ , from same locality, 
taken on various dates early in August”. 
thula^ Colias nastes Hovanitz. [1955, Mar. 9-11 Oct,]’^. Wasmann 
Jour. Biology 13(1): 1-8; figs. 1-2, "These collections [made in 
1952 and 1953] of Colias consisted primarily of the species C. nastes 
(thirty-five specimens) . . "The specimens with which we are 
concerned here were collected at a camp located along this river 
[Meade River, northwestern part of Alaska Territory] at a point 
North 70° 45’ by 156° 30’ West”. "Holotype male and Allotype 
female. Locality: Near the Mead River, Alaska Territory, Alaska, 
N 70° 45’ X 156° 30’ W. Date: Male July 13, 1952, and female 
July 11, 1952 . . . Depository: U . . . S . . . Nat. . . Mus. . Wash- 
ington, D. C. Paratypes: One pair in the author’s collection at the 
California Academy of Sciences, San Francisco, California, One pair 
each in the Canadian National Collection at the Department of 
Agriculture, Ottawa; the American Museum of Natural History, 
New York; and the British Museum of Natural History, London. 
These may be dated 1952 or 1953 . . . The remainder of the series 
are io the collection of the Allan Hancock Foundation”. 

8, COLIAS OCCIDENTALIS SCUDDER. 

alha, Colias dexandra Strecker. 1878 [Sept. - Oct.]"^. Butt. & Moths N. 
Amer., CompL Syo. Catal., p. 81, no. 55a. "... a white $ form 
of rare occurence”. No. locality, series data nor dates given. 
alberta, Colias eurytheme Bowman. 9 Mar, 1942. Can. Ent. 74(2) : 25. 
"Holotype - 6, Wembley, Al[ber]ta,, June 24, 1925, in .the author’s 
collection. Allotype - 2, Wembley, Alta., June 25, 1925, in the 
author’s collection. Paratypes - 7 d , 2 2 , Wembley, Alta., June 
14-25, 1925; 1 6,1 2, Beaver Lodge, Alta., June 29 and July 12, 
1924; 1 6, Fort Vermilion, Alta., June 13, 1925; 1 2, Boswell, 
B. C, July 22, 1927; ^2 2, Rolla, B. C, July 1 and 21, 1927. 
Paratypes in the Canadian National Collection and in the collection 
of the author.” 

dexandra, Colias W. H. Edwards. [1863, Mar.-May - 12 Proc. 

Ent. Soc. Phila. 2(1) : 14-15. Figs. 1-3 on plate 11 may have been 
published later. "From Pike’s Peak; in the Society’s collection; 6 
males, 1 female. The second female is from the collection of Mr. 
George Newman and was taken among the Rocky Mountains . . 

No dates given. 

astraea, Colias W. H. Edwards. Feb. 1872. Trans. Amer. Ent. Soc. 4(?) 
sign. 8: 61. "From a single male taken near the Yellowstone Lake 
by the Hayden Expedition, io 1871”. No date given. 
barbara, Colias Henry Edwards. 5 Feb. 1877’"^. Pac. Coast Lepid. (24) : 
7-8 & 11. "Two females (Coll. Hy. Edw.), Santa Barbara, Baron 
V. Osteo; Gilroy, J. Behrens”. [Santa Barbara & Gilroy are both in 
California]. No dates given. 

Christina, Colias W. H, Edwards. [12 Oct. 1863]’^. Proc. Ent. Soc. 
Phila. 2(2): 79-80. 6 & 2 described. "Taken at the portage of 
Slave River . . .”= No series data nor dates given. 
chrysomelas, C{olias]. Henry Edwards. 5 Feb. 1877’"^ Pac. Coast Lepid. 
(24): 8-9, 11. "Seven 6, ‘five 2. Napa County, Cal[ifornia]. (Coll. 
Hy. Edw.)”. No dates given. Spelled chryomelas by H. Edwards, 
Ibid., p. 9. 

edwmdsii, Colias W. H. Edwards (Behr Ms. name). Jan. 1870. Trans. 
Amer. Ent. Soc. 3(?) sign. 2: 11. "1 6,2 2, from the collection 
of Dr. Behr, taken at Virginia City, [Nevada] at high elevation . . 

No dates given. Spelled edwardsi by Barnes & Benjamin, 8 Dec. 
1926, Bull. Sou. Calif. Acad. Sci. 25(3): 89, under 67a. 


216 


McHENRY 


/. Res. Lcpid. 


emilia, Colias W, H. Edwards. Jan. 1870. Trans. Amer. Ent. Soc 3(?) 
sign. 2: 12. "From 1 6,1 2, taken in Oregon . . . Colleaion of 
Dr. Behr”. No dates given. 

harjofdii, C{olias\ Henry Edwards, 5 Feb. 1877'“^. Pac. Coast Lepid. 
(24) ; 9 & 11. "From seven males, (Coll. Hy. Edw.) Contra Costa 
Co. (Hy. Edw.). Havilah, Kern Co. (R. H. Stretch)”. [Both locali- 
ties in California]. Spelled harfordi by Holland, 1931, Butt. Book, 
rev. ed., p. 294, no. 5. 

hatui, Eurymus alexandra edwardsi Barnes & Benjamin. 8 Dec. 1926. 
Bull. Sou. Calif. Acad. Sci. 25(3): 89, under no. 67a. "Type 
locality; Stockton, Utah. Number and sexes of types: Holotype $, 
VII-30-16; 1 $ Paratype, VII-5-3.” 

krauthii, Colias christina Klots. 24 Jan. 1935. American Mus. Nov. No. 
767: 1-2. "Types. - Male holotype, allotype and thirty-one male and 
seven female oaratypes, from twelve miles west of Custer, Black Hills, 
South Dakota, June 29, 1933; four male paratypes, vicinity of Custer, 
South Dakota, June 27, 1933, and five male and six female para- 
types, Black Idills, South Dakota, July 1, 1931 . . "Holotype, 
allotype, and twelve male and five female paratypes are in the 
American Museum of Natural History; the remainder are in the 
author’s colleaion.” 

lambillioni^ Colias elis Dufrane. 28 Feb. 1947. Bull. & Ann. Soc. Ent. 
Belgique 83(1-2); 70. 2 described. "... Canada, sans nom de 

localite precise . . No date given. 

martini, Eurymus harfordii Gunder. 10 Sept. 1931. Bull. Sou. Calif. 
Acad. Sci. 30(2) : 45. "Holotype 2, . . . South side of Arrowhead 
Lake, San Bernardino Co., Calif. Sept. 2, 1931. Type in Author’s 
coll.” 

occidentalis, Colias Scudder. Sept. 1862. Proc. Boston Soc. Nat. Hist. 
9(?) sign. 7: 109-111; unnumb. fig. (p. 107). "2 6,3 2. Gulf of 
Georgia (A. Agassiz); Fort Simpson, British America. (W. H. 
Edwards,)”. No dates given. 

pallida, C\plias\ alexmidra Cockerell. May 1889. Entomologist 22(311) : 
128, under no. 2. Cites: "... (W. H. Edwards, Can. Ent., 1887, 
229) . . Edwards speaks of two albino females sent by Bruce 
[from Colorado?]. No dates given. 

pallida, C[olias\ christina Cockerell. May 1889. Entomologist 22(311): 
128, under no. 2. Cites: "... (H. H, Lyman, Can. Ent., 1884, 6) 

. . Lyman speaks of two albino specimens of female. No dates 
given. 

palidissima, Colias eurytheme alberta Bowman. 9 Mar. 1942. Can. Ent. 
74(2): 25. "Holotype - 2, Fort Vermilion, Al[ber]ta,, July 15, 

1925, in the author’s collection. Paratypes - 2 2, same locality, 
June 9 and 15, 1925, in the Canadian National Collection and the 
colleaion of the author”. 

shastae, Eurymus occidentalis chrysomelas Barnes & Benjamin. 8 Dec, 

1926 . Bull. Sou. Calif. Acad. Sci. 25(3): 88, under no. 63a. 
"Albinic 2 of chrysomelas . . .”. "Type locality: Shasta Retreat, 
Siskiyou Co., Calif. Numbers and sexes of types: Holotype 6 [in 
error], 1-7 July”. 

weaverae, Eurymus harfordii Gunder. 8 May 1924. Ent. News 35(5): 
156-157; plate 2, fig. G. "Allotype [Holotype] 2, (Author’s Coll.) 
Warner Sorings, San Diego County, California, July 3, 1949”. 

9. COLIAS PALAENO ( LINNAEUS ) . 

chippewa, Colias W. H. Edwards. [Aug. 1870]^'^. Syn. N. Amer. Butt, 
(pp. 7-14) : 85 no. 15. A new name for his own Colias helena. 
"Hab. - Great Slave Lake” [differs from that given for helena\ See 
his helena for data. 


(3):20^-22l , 196} 


NAMES — Pt. 2 


217 


helena, Colias W. H. Edwards. [12 Oct. 1863]^^. Proc. Ent. Soc. Phila, 
2(2) : 80. d & $ described. ''From Mackenzie’s River, . . No 
series data oor dates given. A homonym of Colias helena Herrich- 
Schafifer, 1844’'’. Syst. Bearbeit. Schmett. Europa 1(7): plate 45, 
figs, 206-207 (I have not seen this reference of Herrich-Schaffer) . 

kohlsaati, Eurymus chippewa Gunder. 10 Sept. 1931. Bull. Sou. Calif. 
Acad. Sci. 30(2) : 45. "Holotype . Mt. McKinley Nat. Park, 

Alaska. July 14, 1930”. "Type in Author’s coll.” 

lapponica, Colias palaeno Staudinger. End Jan. 1871. In Staudinger 
& Wocke. Catal. Euron. Faun., p. 5, no. 58a. $ described. Cites 

figs. 403-4 ($, werdandi) on plate 83 in Herrich-Schaffer. 1848’'", 
Syst. Bearbeit. Schmett. Europa 1(36). "Lap. Ross. s.”. No series 
data nor dates given. 

palaeno, Papilio [Heliconius] Linnaeus. 1761 [after 28 July]’®. Fauna 
Svecica. Ed. Alt., p. 272, no. 1014. "Habitat in Pteride rarissime 
Upsaliae, frequentior in Finlandia”. No sex data, series data nor dates 
given. 

philomene, [Papilio'] Hubner. [1805]’“^. Samm. Europ. Schmett. (Papili- 
ones); no text; plate 117, figs. 602-603. Figs. 740-741 on plate 147 
were ptiblished later. 

werdandi. C[olias\ Herrich-Schaffer (Schdnherr Ms. name). 1844’^. Syst. 
Bearbeit. Schmett. Europa 1(3): plate 8, figs. 41-42; 1(7): 102 
(published 1844); plate 83, figs. 403-404 (published 1848). No 
series data, no locality nor dates given. Homonym of Colias 
werdandi Zetterstedt, Insecta Lapponica, page 908, no. 2, [1839]^°. 

10. COLIAS PELIDNE BOISDUVAL & LECONTE. 

interior, Colias Scudder. Sept. 1862. Proc. Boston Soc. Nat. Hist. 9(?) 
sign. 7: 108-109; 1 unnumb, text fig. (p. 107). "5 $, I 2. 
Northern shore of Lake Superior (Prof. Agassiz); mouth Sasketch- 
ewan River, British America (S. H. Scudder.)”. No dates given. 

isni, Eurymus pelidne minisni Barnes & Benjamin. 8 Dec. 1926. Bull. 
Sou. Calif. Acad. Sci. 25(3): 89, under no. 69c. "Type locality: 
Laggan, Al[ber]ta. Number and sexes of types: Holotype $, 16-23 
Aug.” 

lahradorensis, Colias Scudder. Sept. 1862. Proc. Boston Soc. Nat. Hist. 
9(?) sign. 7: 107-108; unnumb. fig. (p. 107). "8 d, 5 2. 
Caribou Island, Strait of Belle Isle, Labrador. (A. S. Packard, Jr.)”. 
No dates given.* 

laurentina, Eurymus philodice Scudder. Mar. 1876. Proc. Boston Soc. 
Nat, Hist. 18(2): sign. 12: 189-190. "... Cape Breton Island” 
[from article title]. "Thirty-nine specimens were collected, of which 
ten were gynandromorphic females, eight pallid females, and the 
rest males”. No dates given. 

minisni, Colias elis Bean. Apr. 1895. Psyche, Jour. Ent. 7(228): 228. 
Intended to withdraw his Ms. name for what he considered the 
albinic female of elis Strecker. 

mira, Colias pelidne Verity. 1905-1911. Rhopal. Palaearctica; p. 347; 
p. xxxiv; plate LXVIII, fig. 49; ol. LXVIII exolanation page, fig. 49. 
"49 ... $ ... (Ravea, Labrador) [e coll. Elwes]”. No date given. 

moeschleri, Colias pelidne Grum-Grshimailo. 30 July 1894. Horae Soc. 
Ent. Rossicae 27 (3-4) : 379, no. 1. 1 2 described. "Specimen 
unum e museo . . . Moeschleri, alterum, in Labrador anno 

1889 collectum, e museo lepidopterologi gemanici Dom. M. Wiskotti 
acceptum.” 

nepi, Eurymus interior Barnes & Benjamin. 8 Dec. 1926. Bull. Sou. 
Calif. Acad. Sci. 25(3) : 89, under no. 65. "Type localitv: Nepigon, 
Ontario. Number and sexes of types: Holotype 2, and 12 Para- 
type both 8-15 July.” 


218 


McHENRY 


/. Res. Lcpicl. 


fieri, Eurymus pelidne skinneri Barnes & Benjamin. 8 Dec. 1926. Bull. 
Sou. Calif. Acad. Sci. 25(3): 89, under no. 69b. "Type locality: 
Yellowstone Park, Wyo[ming]. Number and sexes of types: Holotype 
$ 8-15 July; 2 $ Paratypes, 8-15 July and no date; all being 
original type females of skinneri Barnes.” 
pelidne, C[olias]. Boisduval & LeConte. 1829. Hist. Gen. Icon. Lepid. 
Chen. I’Amer. Sept. 1(7): plate 21, figs. 4-5 (2). Pp. 66-67 
published in livrason 8. 6 & 2 described. "Elle habite le Greenland 
et rislande. M. Sommer, observateur tres-distingue, me I’a aussi 
envoyee comme venant du Labrador”. No series data nor dates given. 
skinneri, Colias pelidne Barnes. 1 Feb. 1897. Can. Ent. 29(2): 41-42. 
"Described from 15 males and 7 females - three of which are yellow, 
three white, and one intermediate — taken in Yellowstone National 
Park, and at Arangie, Idaho, in July”. 

solivaga, Colias W. H. Edwards (Henry Edwards Ms. name). 1877 [Feb. - 
16 Apr.]^’. Trans. Amer. Ent. Soc. 6(?) signs. 1-6: 17, under no. 
68. Gives a Henry Edwards Ms. name as a synonym of interior 
Scudder. 

11. COLIAS PHILODICE GODART. 

alba, Colias philodice Strecker. 1878 [Sept. - Oct.j'^. Butt. & Moths N. 
Amer., Compl. Syn. Catal., p. 82, no. 58a. Cites 2 figures in W. H. 
Edwards, Butt. N. Amer. 2 : plate Colias II, fig. 6 and plate III, figs. 
5-6, 1876. No locality, series data nor dates given here. 
alba, Colias philodice plicaduta F. Chermock. 28 Apr. 1927. Bull. 
Brooklyn Ent. Soc. 22(2): 119. "Holotype 2. July 30, 1924; 2 
paratypes August 17, 1925; 1 paratype August 16, 1925. N. S. 
Pittsburgh, P[ennsylvanila.” 

albida, Eurymus philodice plicaduta F. Chermock. 15 Nov. 1928. Bull. 
Brooklyn Ent. Soc. 23(4): 173. A new name for his own Colias 
philodice plicaduta alba which he considered a homonym of Colias 
philodice alba Strecker, 1878. See his alba for data. 
albinic, Colias philodice W. H. Edwards. [1884, after 12 Sept. - before 
18 Feb. 1885]^^. Trans. Amer. Ent. Soc. 11(3-4): 264, under no. 
68. Cites his own Butt. N. Amer. 2; plate Colias II, 1876. No 
locality, series data nor dates given here. 
anthyale, Zerene Hiibner. 1823 [1 Jan. - 20 Apr.]^^. Ziitr. Samm. Exot 
Schmett. 2nd. 100: 21, no. 154. Figs. 307-308 on plate [54] were 
published earlier but without names. $ described. "Aus Pennsyl- 
vanien”. No series data nor dates given. 
autumnalis, Colias eurytheme eriphyle Cockerell. Mar. 1888 [after 9th]°. 
West Amer. Scientist 4(35): 41-43 (in pt.). "On November 9, 
1887, I caught a female specimen of the autumn brood of eriphyle 
in this locality” [(West Cliff, Custer Co., Colorado)]. 
ehrmanni, Colias philodice plicaduta F. Chermock. 28 Apr. 1927. Bull. 
Brooklyn Ent. Soc. 22(2): 118. "Holotype 2. August 3, 1921. 
N. S. Pittsburgh, P[ennsylvani]a.” 

eriphyle, Colias W. H. Edwards. Apr. 1876. Trans. Amer. Ent. Soc. 
5(?) sign. 26: 202-204. "Some thirty individuals of this species 
and of both sexes, were taken at Lake Labache, in British Columbia, 
. . . , and the whole series came into my possession”. No dates given. 
europome, E\apili6\. D[anaus'). C{andidus\ Haworth. 1803 [in or after 
July]’®. Lepid. Britannica (1): 13, no. 12. "Habitat in Anglia, 
rarissimo. D. Francillon et D. Swainson”. "Descriptio (Ex exemplario 
Germanico)”. No date given. A homonym of europome Esper 
[1777]13, Schmett. 1(1): plate 42, figs. 1-2 (I have not seen 
this reference of Esper) . 

hagenii, Colias W. H. Edwards. [1884, 19 Tan. - 20 Feb.]^"^. Papilio 
3(7-10): 160-161, no. 11; 3(7-10): 163-164. "From 20 6 24 2 
from various localities, from So. Colorado to Montana and Dacotah 


l(}):209-321, 1963 


NAMES ™Pt. 2 


219 


(Bismarck)”. No specific dates given. Spelled hageni by Holland, 
1931 , Butt. Book, rev. ed., p. 297, under 00 . 17. 
hybrida^ Colias philodice Strecker, 1878 [Sept. - Oct.]"^. Butt. & Moths 
N. Amer., CompL Syn. CataL, p. 82, 00 . 58d. 8 cited in W. H. 

Edwards, 1876, Butt, N. Amer. 2: plate Coiias III. "Illinois, 
Georgia”. No date given here. 

invetsata, Colias philodice Nakahara. 27 Jan. 1926. Bull. Brooklyn Ent. 
Soc. 20(5): 223 . "Holotype: $, Midvale, Allegheny Co., P[enn- 
sylvanija., August 20, 1893 . . • Type in the Barnes Collection.” 
kootenai, Colias Cockle. 4 June 191(). Can. Ent. 42(6): 203-204. 
"Kalso and in Northern British Columbia” [from article title]. 
"... May 17 to 20, and fall brood to Oct. 9”. No sex data nor 
series data given. 

laurae, Eurymm eurytheme eriphyle F. Chermock. 9 Mar. 1929. Bull. 
Brooklyn Ent. Soc. 24(1) : 21. 9 described. "Holotype, VIII-1- 

1924: Edmondton, Alberta, Canada.” 

lutetincta, Colias philodice Wolcott. 11 Apr. 1893. Can. Ent. 25(4): 
104, under no. 50. "I first met this form ... in August, 1885, at 
Batavia, Ill'[inoi]s . . . B.ut have since taken it at different times at 
Grand Rapids, [Michigan] . . . and have collected in all about a 
do 2 e,o specimens, one of them a female”. 
maria, C{olias\ philodice W. H. Edwards (Grote Ms. name). 13 July 
1885. Papilio 4(9-10): 171. Inadvertently describes a cyanide 
stained specimen. 

melanic, Colias philodice W. H. Edv/ards. [1884, after 12 Sept. - before 18 
Feb. 1885]^^. Trans. Amer. Ent. Soc. 11(3-4): 264, under no. 68. 
Cites [$] in his Butt. N. Amer. 2: plate Colias III, 1876. No locality, 
series data nor dates given here. 

minor, Colias philodice plicaduta F. Chermock. 28 Feb. 1927. Bull. 
Brooklyn Ent. Soc. 22(2): 118. "Holotype $. June 28, 1925. 
N. S. Pittsburgh, P[ennsylvanPa.” 

miscidke, E[urymm]. p\hilodice\ Scudder. 1 June 1889. Butt. East. U. S. 
& Canada 2(8): 1115. "Mr. F. G. Sanborn has shown me a very 
interesting suffused female of this species taken by Mr. John Osgood 
in Lynn, Mass[achusetts]., in August, 1863’h 
nig, Colias philodice Strecker. 1878 [Sept. - Oct.]"^. Butt. & Moths N. 
Amer., CompL Syn. CataL, p. 82, no. 58b. S from Montreal, Canada 
described; also cites W. H. Edwards, 1876, Butt. N. Amer. 2: plate 
Colias III, figs. 8-9 (copied from Glover’s unpublished plate). No 
dates given. 

nigficosta, Eurymus eurytheme eriphyle F. Chermock. 9 Mar. 1929. Bull. 
Brooklyn Ent. Soc. 24(1): 21. " . . . occurs in the female sex 
only”. "Holotype, VIII-1-1924; Topoparatype, VIII- 1-1 924; Ed- 
mondton, Alberta, Canada”. 

nigridice, Eurymus phil[odice]. Scudder. 1 June 1889. Butt. East. U. S. 
& Canada 2(8): 1114. d & $ described. Cites W. H. Edwards, 
1876, Butt. N. Amer. 2: plate Colias III, figs. 8-9 (S ). Cites fig. 5 
on Glover’s unpublished plate I. Also cites other specimens which 
were not in his possession (one captured 29 July 1883). 
nigrina, Colias philodice Strecker. 1900 [Mar. 9th - 13th]^. Lepid., 
Rhopal. & Heter., Indig. & Exot. SuppL No. 3: 19. Applies the 
name to what he formerly called nig. Here he cites 3 specimens: 
"... Bethlehem, P[ennsylvanPa. . . . Montreal, Cao[ada] . . . Orillia, 
Can.”. No dates given. 

nigrofasciam, Colias philodice Reiff. 15 Jan.- 1917. The Lepidopterist, 
Off. Bull. Bost. Ent. Club. 1(3): 22-23; plate 3, 1 unnumb. fig. 
2 described. "... was caught . . . during August, 1916 in Massa- 
chusetts, and it is now in the collection of the author”. 


220 


McHENRY 


/. Res. Lcpid. 


notatus, Papilio Megerle. 28 Nov. 1803. Catalogus Insectorum Qua 
Viennae Austriae 28. Novembris 1803 Auctionis Lege Distrahuntur, 
p. ?, no. 436. "M. ex. Georg[ia] . . . 6’’. No dates given. I have 
not seen this reference, all data is from Clark & Clark. 
notatus, Papilio Clark & Clark (Megerle Ms. name). 30 Apr. 1941. 
■Proc. Ent. Soc. Washington 43(4) : 81, no. 436; & page 83, no. 1. 
"M. ex Georg[ial. . . 6”. No dates given. Reprint Megerle’s 
description. 

pallidice, E{urymus\ p[hilodice\. Scudder. 1 June 1889. Butt. East. U. S. 
&c Canada 2(8) : 1115; pi. 7, fig. 6; pi. 7 explanation page, fig. 6. 
Page 1125 (published later). $ described. Cites W. H. Edwards, 
1876, Butt. N. Amer. 2: plate Colias II, fig. 6 and plate Colias III, 
figs. 5-6. Also cites fig. 11 on Glover’s unpublished plate 6. Data 
is general. 

philodice, Colias Godart. 1819. In Godart & Latreille in Latreille. Encycl. 
Meth. 9(1) : 87, no. 35 (named in French); 9(1) : 100-101, no. 35. 
[6] & $ described. "File se trouve dans I’Amerique septrentionale, 
particulierement dans la Virginie”. No series data no dates given. 
Spelled philodin by W. H. Edwards, 1872, In Hayden, Prelim. Rept. 
U. S. Geol. Surv. Mont. & Port. Adj. Terr. 5th. Rept. Progress, p. 466. 
plicaduta, Colias philodice Nakahara. 27 Jan. 1926. Bull. Brooklyn Ent. 
Soc. 20(5) : 222-223. "Holotype: $, Ithaca, N. Y., July 30, 1923 
. , . Paratopotype : $, July 16, 1924. Paratype: $, Lava, Sullivan 
Co., N. Y., "June.” Holotype has been presented to the Barnes 
Collection, Decatur, Illinois]., and paratype is in the same collection. 
Paratopotype is retained in the collection of the writer”. 
raritus, Eurymus philodice Gunder. 30 July 1928. Can. Ent. 60(7): 
163; plate A, figs. 2, 2a. "Data: Holotype $ . . . Scranton, 
P[ennsylvani]a. (bred . . .), July 8, 1927. In the Author’s coll.” 
reducta, C\qlias\ philodice anthyale Dufrane. 28 Feb. 1947. Bull & Ann. 
Soc. Ent. Belgique 83(1-2): 69. $ described. ”. . . Temple, 

P[ennsylvani]a. . . ”. No series data nor dates given. 
fothkei, Colias philodice Reiff. 15 Aug. 1917. The Lepidopterist, Off. 
Bull. Bost. Ent. Club. 1(11): 84; plate 7, 1 unnumb. fig. "Col- 
lected August 27, 1905 in the Susquehana Valley, P[ennsylvani]a., 

. . . Type 1 male in Mr. Rothke’s collection”. Mentions another 
"specimen (no status). 

s errata, Eurymus philodice F. Chermock. 9 Mar. 1929. Bull. Brooklyn 
Ent. Soc. 24(1): 21. "... occurs in tlie male sex only . . .”. 
"Holotype, VIII-8-1926; paratypes. No. 1, VIII-8-1926; Nos. 2, 3, 4, 
VII-10-1926; No. 5, VIII-12-1926; Rossgrove, near Aspinwall, 
P[ennsylvani]a.” 

suffusa, C[olias]. philodice Cockerell. Mar. 1889. Entomologist 22(310): 
55. Cites: ". . . Massachusetts, Maynard, Butt, of New Eng., pi. VII, 
fig. 57c”. No date given. 

virida, Colias philodice Strecker. 1878 [Sept. - Oa.]"^. Butt. & Moths N. 
America, Compl. Syn. Cat., p. 82, no. 58c. $ described. "One 

example . . Type locality: [near Montreal, Canada]? No date 
given. 

vitahunda, Colias chrysotheme Hovanitz. 30 July 1943. American Mus. 
Nov. No. 1240: 2, 3-4. "Holotype Female and Alotype Male. — 
McKinley National Park, Alaska, July 18 to August 9, 1930, . . ., 
J. D. Gunder Collection, in the American Museum Natural History. 
Paratypes. — Twelve males same data as above; eleven males same 
locality but July 29, 1931, . . .; for males July 15-20, 1931, ... in 
Los Angeles Museum; ten while female same data as holotype; four 
white females July 20-30, 1931, ... in Los Angeles Museum; one 
white female August 9, 1930, ... in Los Angeles Museum”. 


l{})X20^-22l , 196} 


NAMES — Pt. 2 


221 


12. COLIAS SCUDDERII REAKIRT. 

flavotincta, Colias scudderi Cockerell. Apr. 1901. Psyche, Jour. Ent. 
9(300) : 186. Cites W. H. Edwards, Butt. N. Amer. [1]: plate 10, 
fig. 5 (?) (part of illustrated series of scudderii, 1872). 
ruckesi, Colias scudderi Klots. 2 Nov. 1937. Jour. New York Ent. 
Soc. 45(3-4): 324-326. "Holotype male, allotype female, nineteen 
male and one female paratypes, from Windsor Creek Canyon, west 
of Cowles, N. Mex., July 2, 1935, . . . Twenty-five male and nine 
female paratypes from the same locality, July 4, 1936, . . . All were 
taken in about middle Canadian Zone, at from 9000 to 9500 ft. 
altitude, in grassy meadows surrounded by forest, along the Forest 
Service trail about halfway between Cowles and the summit of 
Santa Fe Baldy Peak. Holotype, allotype, six male and two female 
paratypes deposited in the American Museum of Natural History; 
four male paratypes deposited in the U. S. National Museum; four 
male paratypes deposited in the Canadian National collection; the 
remainder of the paratypes at present in the author’s collection.” 
scudderii, Colias Reakirt. [12 June 1865]^^. Proc. Ent. Soc. Phila. 4 
(pp. 213-339) : 217-218, no. 2. "Hab. — Rocky Mountains, Colo- 
rado Territory. (In my collection, 8, and that of the Entomological 
Society. $.)”. "This species was collected in August, 1864 . . .”. 
Spelled scudderi by Holland, 1931, Butt. Book, rev. ed., p. 295, 
no. 10. 


FOOTNOTES 

IVol. 6 title page date is qu*Iified by indirect date data (certain names in Fabricius’ article 
were mentioned in the Allgemeine Literatur-Zeitung, Halle [Rnal 2(303): cols. 1177-1181, 19 
Dec. 1807). 

2Pt. 2 title page date is qualified by indirect date data by A. S. Corbet, 28 Feb. 1939, Jour. So. 
Bibliop-. Nat. Hist. 1(7): 96. 

^Hemming. 1937. The Generic Names of the Holarctic Butterflies 1: 139, see no. 376. 

"^Work title page date is qualified by indirect date data on pages 39 & 48 in the Sept. & Oct. 

numbers of Vol. 3 of the Bull. Brooklyn Ent. Soc. 

5Dos Passos. 1962. Jour. Lepid. Soc. 16(1): 45-46. 

^Sherborn & Woodward. 1901. Ann. & Mag. Nat. Hist. Ser. 7. 7: 137-140. 

7No. 3 title page date is qualified by the preface date and a Ms. date of receipt (13 Mar. 1900) 
written on a copy of No. 3 in the library of the Los Angeles County Museum. 

®No. 35 title page date is qualified by the article signature date. 

9The date Sept. 1869 (given for Pt. 4 as per footnote 16) is qualified by the receipt date or 
Pt. 4 (see Trans. Amer. Ent. Soc. 2; XVIII, 1868-9). 

’OTome 5 title page date is qualified by a meeting date (see p. 383). 

^ IKirby. 1871. Syn. Catal. Diurn. Lepid., p. 494, no. 29a. 

1 ^Article received by publisher 9 Mar. 1955. No. 1 containing the article was received at the 
Allen Hancock Foundation Library (Univ. Sou. Calif.): 11 Oct. 1955. 

l^No. 1 title page date: Mar., Apr. & May 1863. No. 2 receipt date (see Proc. Ent. Soc. Phila., 
2: 164). 

14More actual publication date data is needed. 

'5Proc. Ent. Soc. Phila. 2: 164 (No. 2 receipt date is given). No. 2: pp. 61-162. 

1 ^Hemming. 1931. Proc. Ent. Soc. Lond. Ser. A. 6: 42-45. Pp. 7-14 of Synopsis were published 
with Butt, N. Amer., pt. 6. 

17Hemming. 1937. Hubner 1: 579-589. 

1 ®Title page date is qualified by the preface date. 

^ ^Hemming. 1937. Hubner l:146-3Z‘t. 

20Sherborn, 1922. Index Animalium, 2nd. Ser. Vol. A-B: cxxxi. 

21 Vol. 6 title page date is qualified by the date of signature 3 and the date of a letter of W. H. 
Edwards to H. Edwards (Henry Edwards correspondence in the Amer. Mus. Nat. Hist, N. Y.) 
dated 16 Apr. 1877 and stating he would mail a copy of the catalogue the next day. 

22No. 2 (Vol. 11) of the Trans. Amer. Ent. Soc. was available 12 Sept. 1884 (see p. xxxvi, 

vol. 11). The issue date of the separate was 18 Feb. 1885 (see page 2 of separate). 

23Hemming. 1937. Hubner 1: 451-487. 

24Nos, 7-10 (v. 3) were not published before 19 Jan. 1884 (see indirect date data on page 193). 

No, 1 (v. 4) was published 20 Feb. 1884 (see p. 42, v. 4). 

25pp, 221-330 (Mar. & Apr. 1865) receipt date (see Proc. Ent. Soc. Phila., v. 4, p. xii). 










y 




tMt. 



j r n 
>■: 


,1 


.j4. 


>- ' .f» 1-1 

• . 'AV* , 


r 




Journal of Research on the Lepidoptera 1 (3) :223-235, 1963 

1140 W. Orange Grove Ave., Arcadia, California, V.S.A. 

© Copyright 


A STANDARD METHOD FOR MOUNTING 
WHOLE ADULT LEPIDOPTERA ON SLIDES 
UTILIZING POLYSTYRENE PLASTIC 

CHARLES L. HOGUE 

Los Angeles County Museum, Los Angeles 1 , California 

Comparative morphological studies of insects are greatly 
facilitated if the material is prepared and mounted in a way that permits 
its easy manipulation and rapid, yet detailed, examination. Mounting 
specimens dry, on pins, has been the method of preference of Lepid- 
opterists virtually exclusively, but it does not permit detailed examina- 
tion of integumentary features that are of great taxonomic importance. 
Attempts to interpret the relationships of higher categories in the 
lepidoptera entirely on the basis of structure externally visible on such 
specimens have, in many cases, led to errors and have confused the 
taxonomy of many groups. 

Although most workers are willing to utilize slide mounts for limited 
parts of the body, especially the genitalia, they have not extended the 
treatment to the whole animal and prefer to keep their material in 
fluids, usually alcohol. While alcoholic specimens allow the greatest 
amount of freedom of examination, considerable time and patience 
is spent simply handling them — ^unstoppering vials, pouring alcohol, 
restoppering vials, etc. As a result, a worker may hesitate to make the 
innumerable comparisons and recomparisons requisite to phylogenetic 
analysis. Furthermore, since the specimen must be examined with 
relatively low power dissecting microscopes, many fine details, such as 
microchaetae and other minute sense organs, surface textures, striae, 
spicules, spiracular structure, etc., are not visible, or at least not clearly 
observable. Among the other disadvantages of alcoholic specimens are* 
that they do not keep stains well, are subject to evaporation and destruc- 
tion from drying and suffer damage due to repeated handling. 

Slide mounts, on the other hand, permit very detailed observation 
and are very easily handled. Admittedly, they allow the specimen to 
be viewed only from a limited number of aspects, but if prepared in 
a careful manner, such as that described below, most of the integu- 
mentary anatomy is available for inspection. Slides may be stacked as 
high as 4 or 5 deep and the specimens readily compared by focusing 
up and down with a dissecting microscope or detailed drawings rapidly 
prepared for the same purpose with a microprojector. In addition, they 
may be permanently stained, are subject to destruction only by breaking, 


223 


224 


HOGUE 


/. Rex. Lepid. 


and may be handled indefinitely without wear. Even fairly large Lepid- 
optera (e.g., smaller Catocala, Vanessa) may be placed on slides but I 
do not recommend it for very large species (e.g., saturniids, sphingids). 

The details of the method outlined below were worked out on 
noctuid moths of medium size with relatively simple genitalia. Yet I 
believe that most groups of Lepidoptera may be similarly treated, 
although, of course, certain groups may require special handling be- 
cause of extremely small size or peculiar modifications, especially in 
the genitalia. 

The general treatment below might be applied advantageously to 
many insect orders (especially Neuroptera, Trichoptera, Diptera, etc.). 
All stages of mosquitoes are beautifully accommodated and I have had 
promising results with Collembola, a group notoriously difficult to 
mount on slides. The latter usually shrivel badly in balsam and eventu- 
ally deteriorate if placed in aqueous media, apart from being flattened 
or otherwise distorted. 

THE USE OF POLYSTYRENE PLASTIC 

Whole mounts of most insects require fairly thick preparations. 
Workers who have tried to make such preparations with the usual 
natural (Canada balsam, Euparal), synthetic (Permount, Diaphane), 
or mixed aqueous (PVA, Hoyer’s) media find that there are certain 
difficulties: These media, when applied in large amounts take a very 
long time to -harden, specimens invariably move in them from their 
desired positions during the hardening period (even if the laborious 
and time consuming "layering” or "build-up” technique is used), and 
they change color or deteriorate after a time. 

Polystyrene plastic' (hereafter referred to simply as "plastic”), 
however, primarily because of its setting qualities, is the ideal medium 
for holding dissected parts in their desired positions on slides. Because 
it gels evenly and rapidly throughout, any specimen, regardless of shape 
or center of gravity, may be orientated perfectly in any position. Further 
advantages of plastic over the media mentioned above are its greater 
transparency and resistance to discoloring, and its thorough hardness 
and permanency when finally cured. In addition, it has the desired 
qualities of being its own clearing agent and of slightly intensifying 
certain stains. It has one disadvantage at present; it cannot be dissolved 
gradually and without fragmenting after complete hardening by any 
agent I know and therefore a specimen cannot be removed once 
mounted. This is a serious drawback, however, only with scarce material 
which might need to be remounted for study from a different aspect. 
The methods described in this paper are intended primarily for morpho- 
logical and phylogenetic studies where abundant material is available. 

In any case, since slides are rarely remade unless very poorly prepared 


'Known commercially by many trade names, this is the type used for embedding biological 
specimens and other items by hobbyists. 


( 3 j ; 225 - 2 j 5 , 196} 


PLASTIC METHOD 


225 


in the first place, the insolubility of plastic should not limit its use. 
Preparations on broken slides may be soaked off in cellosolve and glued 
onto new slides with a thin layer of plastic. 

The steps involved in using plastic as a medium for thick micro- 
scope slide mounts are as follows: 

1. The specimen, after dissection and dehydration in up to 95% ethanol 
(see directions below), is transferred to a mixture of 50% acetone and 50% 
uncataly 2 ed plastic (because of the difference in density between these compon- 
ents they do not mix readily- — -mixing may be accomplished by shaking them 
together in a vial for a few seconds). Since the acetone evaporates rapidly 
from the mixture, its container must be kept covered as much as possible. The 
mixture clears and completely dehydrates the specimen; the specimen should 
remain in it about fifteen minutes. 

2. While the specimen is clearing, a slide is prepared to receive it in the 
following manner: Uncatalyzed plastic is dropped onto the slide and spread 
out over an area the size of the cover slip. Spreading is done with a glass rod 
(3mm. diameter), which has been previously dipped into catalyst. The rod is 
used to apply and thoroughly mix the catalyst into the plastic. The depth to 
which the rod is dipped determines the amount of catalyst picked up. I find 
that dipping it about 5mm. provides the right amount of catalyst to gel the 
thickest layer of plastic that can. be spread over a 22mm. square area (size 
of standard, square cover slip) without running over. Enough plastic should 
be applied to just cover the thickest part of the specimen. If the specimen is 
very thick and the viscosity of the plastic is not enough for it to stand level 
with the specimen, more layers must be added. 

3. The specimen is next transferred to the plastic on the slide and 
generally oriented. 

4. The slide is then placed on a warming plate, previously heated to 
about 130°F. to accelerate the gelling of the plastic. While the preparation is 
on the plate, it is viewed with the low power of a dissecting microscope and 
the specimen appropriately oriented. The specimen is held in its desired posi- 
tion with dissecting needles until the plastic gels enough to firmly anchor it. 
At this temperature, gelling will begin rapidly and will be far enough advanced 
to hold the specimen in about 7-10 minutes^. Impending gelation can be noted 
by the cessation of movement in the plastic revealed by dust particles present 
in it. 

5. The slide is allowed to remain on the warming plate about 15 minutes 
to ensure thorough gelation. 

6. The preparation is then ready to receive a cover slip if the first layer 
of plastic was sufficient to cover the specimen. If not, more layers must be 
added in the same manner as the first. 

7. Glass chips or other cover slip supports are unnecessary and should 
not be used since they hinder the settling of the cover slip on the slightly 
shrinking plastic, and may cause a broken preparation » 

8. The cover slip is prepared by covering it with just enough catalyzed 
plastic (applied with glass rod as above) to spread over its entire area when 
inverted and placed on the specimen. When the cover slip is in place the 
whole preparation is returned to the warming plate for gelling. The heat also 
helps the plastic under the cover slip to spread. If too little plastic was applied, 
additional amounts may be added at the side of the cover slip until all the 
spaces are filled. Any excess plastic may be wiped off from around the edges 
or, if gelling has occurred, trimmed with a blade. 

9 . Complete hardening and curing of the whole preparation can be 
accomplished by leaving it overnight in an oven at about 100°F. or can be 
allowed to take place slowly at room temeprature. 

2This figure varies according to the formulation and age of the plastic. I find these times average 
but they may be longer especially for formulations designed for long shelf life. 


226 


HOGUE 


/. Res. Lepid. 


STANDARD DISSECTION AND MOUNTING 


Four slides are required to hold the various parts of the whole insect 
(fig. 1): 1. The main part of the body including the two halves of 
the head, thorax and abdomen are put on the first slide (the tegulae, 
antennae and basal portions of the right wings being dissected away— - 
see below) (fig. 1:1); 2. A second slide holds the legs, basal portions 
of the right wings, antennae and tegulae (fig. 1:2); 3. The wings 
occupy a third slide ( fig. 1:3); 4. The genitalia are put on a final 
slide (be they male (fig. 4b) or female (fig. 4a). 

These main parts are dissected and mounted as follows: 

Preliminary general dissection 

The following instructions apply to specimens mounted on pins. Specimens 
in alcohol or otherwise preserved are excused from the impertinent steps. 

1. The left pair of wings are broken off at their bases and the right pair 
cut off transversely with scissors beyond the end of the frenulum. The wings, 
with the data labels removed from the pin and a slide reference label, are set 
aside in a cellophane envelope. 

2. The body of the insect, still on its pin, is wetted in 75% ethanol 
and then boiled in water for 5 minutes. This softens it and allows the pin to 
be removed easily. 

3. The body is next transferred to a solution of 10% potassium hydrox- 
ide which is heated to near boiling in a small beaker or test tube suspended in 
a water bath. The specimen is left in the solution until thoroughly macerated 
( 15 minutes) . 

4. After maceration, the body is transferred for dissection and cleaning 
to distilled water in a watch glass, preferably one with a slightly convex 
bottom. 

5. Next the genitalia are removed. In the male this is done by severing 
the membrane between abdominal segments VII and VIII (or further cephalad 
to include special modifications on more proximal abdominal segments if 
present). The female is treated likewise, but by severing the membrane between 
segments VI and VII. The genitalia are rubbed clean of scales (I like to use 
a pointed applicator stick for this; others may prefer a fine camel’s hair brush 
or like instrument) and transferred to clean distilled water. 

6. The legs are now removed (coxae and trochanters should remain 
intact), rubbed clean of scales and transferred to clean water. 

7. The body itself (head, thorax with bases of legs, and abdomen minus 
genitalia) is finally bisected with a sharp, single-edged razor blade. Great care 
must be taken at this step to insure as perfect a bisection as possible. The 
specimen is held ventral side up in the watch glass and a mid-ventral, longi- 
tudinal slit is made at the posterior end of the abdomen with scissors. The 
razor blade is then brought down on the mid-ventral line guided by this slit, 
the coxae and the two halves of the proboscis between which its edge is 
placed. The blade is first used to compress the body gently and then pressed 
firmly against the bottom of the watch glass and rocked to and fro to make 
the final cut. The slightly convex bottom of the watch glass helps to make a 
clean cut. After the halves of the body are separated, cleaned and rubbed free 
of scales, they join the other parts in distilled water. 

8. Dehydration in 75% and 95% ethanol follows (if no aqueous stain 
is used— -see "STAINING”). The female genitalia and phallus of the male 
genitalia go dirertly to 95% ethanol to harden their membranous parts. This 
is done in a special manner as described below^ 


fj).- 223-23 5, 1^63 


PLASTIC METHOD 


227 


I Ai, I 

-. -\Ll.. ^•?si4 r«F, ^ ^ 


-X r ^ ^^ .. 

!,„ ^--'x .) 


v'‘ J*-"' 






<v/ r 




CiJ?' f’lTA A- 



g/i^ ahia S01 

CM/: 

- £>. 

mMB,. 

ca^\^'Z03J/-'j 


3 


i 




Sw 

i?kf. Cijfi^m’ 

C Twft'ifcfe&j 


I<.#s4 «SfL 

CLnm0s/~f 


4 a 




4f?f4 d/ii^a Sm. 


a/^efm4 /# 


db 


Fig. 1: Set of slides (I - 4) holding a whole specimen {4a example 
a female; 4h a male) . 


228 


HOGUE 


/. Res. Lepid. 



(3):22}-2}5, 1^63 


PLASTIC METHOD 


229 



Figs. 2-6: Close-up views of slides shown in fig. 1 to show detail of 

arrangement of parts. Fig. 2: Main body, bisected. Fig. 3: Miscellaneous 
parts. Fig. 4: Wings. Fig. 5: Female genitalia. Fig. 6: Male genitalfa. 


230 


HOGUE 


/. Res. Lcpid. 


9. The wings, previously set aside in a cellophane envelope, are bleached 
for about 2 or 3 minutes (at least long enough for the veins to become well 
defined) in 5% sodium hypochlorite (full strength household bleach — Purex, 
Chlorox), washed in distilled water, and dehydrated by passage through 75% 
and 95% ethanol. 

If it is desired that the veins be stained a totally different procedure 
is necessary: 

After wetting in ethanol and soaking for 20-30 minutes in hydrogen 
peroxide ( 3 % ) , the scales are carefully rubbed off with a blunt camel’s hair 
brush. The denuded wings are then washed in distilled water and transferred 
to the stain. A long destaining period must follow in order to decolorize the 
membranes. 

The dissected parts, now in 95% ethanol, are ready for fine, detailed 
dissection and cleaning (and staining with non-aqueous stains if desired — see 
"STAINING”) before mounting in plastic on slides. I find that these two 
jobs are best done in the 95% ethanol; since the specimen has become fairly 
brittle, sclerites are easily separated and remaining unwanted scales easily 
flaked off. 

Final dissection and mounting 

The different parts are treated in various ways. Reference to the 
figures will clarify the descriptions (including terminology) of the fine 
points of dissection and orientation on the slide. 

Slide 1: Main body (figs. 1:1;2) 

The two halves of the main body of the specimen require only 
minor alteration before being transferred to the clearing mixture. The 
tegulae and antennae are removed and the bases of the right pair of 
wings are carefully dissected away with the axillary sclerites. These 
parts are mounted with the legs (see "Slide 2”). If the specimen is a 
noctuoid moth, the ventral portion of the second phragma is also cut 
away with fine scissors to allow an unobstructed inner view of the 
tympanal area. Similar special treatment may be necessary with other 
taxa. 

The halves are mounted on the slide, one with its outer surface up 
and the other (the one with the cut phragma if noctuoid) with its 
inner surface up. 

Slide 2: Legs, etc. (figs. 1:2;3) 

The legs, bases of wings (with axillary sclerites), tegulae and 
antennae are all mounted together on the same slide since they are of 
about the same thickness and do not orient well if left attached to the 
main body. Usually these parts require no further dissection although 
sometimes one may desire to separate the components of the pretarsus 
or the antennae. Their orientation is best explained by the figures. 

Slides: Wings (figs. 1:3;4) 

No special treatment is needed for the wings. After dehydration 
they are simply mounted flat in the plastic ( which also clears them ) . 
If too large to fit on a standard microscope slide, they may be accom- 
modated by a large sized .slide (2"x3^')- if very large, they may have 
to be cut up into sections. 


if3j-"225-2j5, 1963 


PLASTIC METHOD 


231 


Slide 4: Genitalia — female (figs. l:4a;5;7) 

The following procedure is considerably modified from that usually 
employed by lepidopterists. The conventional procedure is to leave 
abdominal segment VII attached and to mount the entire undissected 
complex ventral side up or laterally on the slide. This, however, has 
the following undesirable features: (1) segment VII, still attached, 
obscures detailed viewing of the structures beneath, (2) if mounted 
with the ventral side up the venter of segment VIII is plainly visible 
but the pleura, which also may possess important characters, are not. 
Furthermore, only the ventral edges of the ovipositor lobes are in plain 
view; their profile, patterns of sclerotization and chaetotaxy are impos- 
sible to see. If mounted laterally, the reverse viewing difficulties are 
present. (3) The whole complex is cylindrical in general shape and 
rolls easily, making it hard to position in a standard way. 

By mounting the female genitalia as described below, these diffi- 
culties are surmounted. The steps involved are as follows (fig. 7 shows 
them partly completed) : 

1. The genitalia have been freed from the remainder of the abdomen 
and cleaned of most of their scales (see above, "Preliminary general dissec- 
tion”)- 

2. Segment VII is removed by splitting it laterally and tearing it off. The 
tear should follow the line of the membrane between it and the following 
segment. Care must be taken not to damage the sclerites around the ostium 
bursae. 



Fig. 7 : Oblique dorsal view of female genitalia showing principal 
structures in state of partial dissection (diagrammatic). 


232 


HOGUE 


/. Res. Lepid. 


3. Next, the genitalia proper are slit mid-dorsally, a cut being made 
first with scissors through segment VIII and continued by tearing caudad to the 
tip between the ovipositor lobes. 

4 . The genitalia are then opened up, the oviduct and rectum are pulled 
up and cut free, and' the whole complex cleaned. 

5. The bursa copulatrix is next inflated with water in much the same 
manner as the vesica of the male phallus (see below). The cannula is inserted 
into the ostium bursae and passed down the ductus bursae. The stream of 
water from the syringe cleans the corpus bursae internally and expands it 
into its full shape. Sometimes a tiny hole is needed to allow the escape of the 
fluid and debris. It is usually not necessary to go through a hardening process 
with the bursa copulatrix as is the case with the male phallus vesica since it 
generally holds its shape better than the latter, 

6. Staining with aqueous stains follows if desired. 

7. Preparatory to dehydration in 95% ethanol the genitalia are opened 
again and placed flat (inner surface down) in a stender. A small piece of 
glass (broken microscope slide) is placed over the opened portion (not over 
corpus bursae! ) to hold it flat while the water is withdrawn from the vessel 
and replaced with 95% ethanol. The same may be done for segment VII, 
previously removed. It may be necessary, if not already done, to punch a small 
hole in the corpus bursae before adding the ethanol if the ductus bursae is 
so narrow that it would prevent the free exchange of liquids. Otherwise, the 
diffusion gradient — ethanol diffusing out faster than water diffusing in — ^may 
cause the collapse of the bursa. 

8. Final cleaning may be carried out after dehydration. The preparation 
is then ready for clearing and mounting. 

Slide 4: Genitalia — male (figs. 1:4b; 6; 8-10) 

Lepidopterists disagree on dissection procedures and orientation 
of the male genitalia on slides. The majority prefer to leave all the 
elements intact and mount the complex ventrally with the valves spread 
open as widely as possible, the least troublesome method. Others also 
do not separate the parts but mount the whole complex laterally. Still 
others dissect various parts away, often only the phallus whose vesica 
may or may not be inflated and the main orientation is either ventral 
or lateral. 

These variations may be dictated by either the whims of the investi- 
gator or the nature of the material, i.e., the positions of the characters 
of greatest significance. Too often it is the former. Obviously, the only 
scientific and rational approach to the problem is to dissect and mount 
many specimens in many ways, even keeping some unmounted in 
fluids. Yet when time is limited, and many comparisons are to be made, 
a single, standard method must be chosen. For this, I prefer to mount 
the main structure of the genitalia laterally, with one valve and other 
elements such as the juxta, anellus, etc. removed and mounted separately. 
Furthermore, I always inflate the vesica of the phallus; though this is a 
time consuming task, the results are well worth it. 

Even apart from the conditions just mentioned, I believe that this 
procedure is superior to the undissected ventral mount for morpholo- 
gical study since there are generally better, undistorted viewing aspects 
for the majority of characters and additional characters are revealed. 
Specifically, (1) the uncus is natural in shape and position instead of 


i(}):22}-2}5, 196} 


PLASTIC METHOD 


233 


being folded or angled if long, or viewed on end if short. Its entire 
profile is observable in plane view; (2) the tegumen is untwisted 
( otherwise usually resulting from the disturbed uncus ) ; ( 3 ) the overall 
shape, lobes and spines etc. of the vesica are fully exposed, not com- 
pressed into the body of the phallus; (4) the anellus is not obscured 
by the juxta (and phallus too if left intact^; (5) the valves (at least 
the one dissected away and usually also the one intact) lie completely 
flat so that their median structures are not distorted by foreshortening 
as is usually the case in a ventral mount. 

The steps followed in dissecting the male genitalia are as follows: 

1. The genitalia have been freed from the remainder of the abdomen 
and cleaned of most of their scales (see above, ''Preliminary general dissection”). 

1 2. The phallus is removed by grasping it at the tip, i.e., posterior end 

protruding between juxta and anellus (fig. 8), and pulling gently to the rear 
(fig. 9). If the phallus does pot so protrude, the forceps may need to be 
inserted a short distance into the canal inclosing it to gain a hold. If the 
phallus is withdrawn properly and carefully, the manica will turn wrong side 
out and the ductus ejaculatorius will pull out through the hole in it (fig. 9). 

If staining with aqueous stains is desired at this point, the main portion 

of the genitalia should be set aside in distilled water. Otherwise, they may go 

directly into 95% ethanol. 

3 . The ductus ejaculatorius should now be snipped off with a pair of 
fine scissors a short distance from the aedeagus, 

4. Next comes the preliminary inflation of the vesica. This is accom- 

plished by first teasing as much as possible of the vesica out of the aedeagus 
with fine forceps and then inflating it with a hypodermic syringe filled with 
distilled water. I find that a 2cc. syringe tipped with a glass cannula (in 

preference to a regular hypodermic needle which does not permit the variety 

of sizes needed) best for this work. The cannula is inserted into the ductus 
ejaculatorius (fig. 10) and pressure applied until the vesica completely inflates 
and everts. It is usually necessary to crimp the ductus with forceps against the 
cannula at the point of insertion to prevent leakage and the phallus from 
coming free under the hydraulic pressure. 

5. The vesica is next hardened to preserve its fully inflated shape (fig. 
10). This step follows staining if aqueous stains are used. Several steps are 
necessary in this process: 

a. The phallus is placed in clean distilled water in a watch glass. 

b. A short length ( 1 cm. ) of silk surgical suture ( cardiovascular 
5-0 or 6-0) is tied into a loose, simple knot and also put into the watch glass. 

c. The hypodermic syringe is filled with 95% ethanol. 

d. The cannula is inserted as above. This may be difficult since the 
alcohol rapidly diffuses out of the tip setting up a strong current which 
constantly carries the phallus away, 

e. After the cannula is properly inserted the phallus is tied to it 
with the suturing thread (around the center of the aedeagus) by slipping the 
loose knot over it and drawing it tight with forceps (fig. 10). Some kind of 
clamp or modelling clay is useful at this point to hold the syringe while both 
hands are being used to tie the knot, 

f. The water in the watch glass is now withdrawn and quickly 
replaced with 95% ethanol. 

g. Pressure is exerted with the syringe and maintained for a short 
while (ab^out 3 minutes usually suffices) until the vesica is thoroughly 
dehydrated and hardened. Leakage of the ethanol from the end of the vesica 
is prevented by clamping it with fine forceps (fig. 10), 

h. The phallus is slipped off the cannula and the suturing thread 
untied and removed. 

i. The phallus joins the rest of the genitalia in 95% ethanol. 


234 


HOGUE 


/. Res. Lepid. 



8 



Figs. 8-9: Ventral views of male genitalia showing principal parts and 
procedure for removal of phallus. Insert showing relationship^ of components 
of phallus complex in lateral, sectional view. (All figures diagrammatic). 


( 3 j .- 223 " 2 j 5 , 1963 


PLASTIC METHOD 


235 


10 



Fig. 10: Phallus rf male genitalia properly prepared for inflation and 
hardening of ¥esica (diagrammatic). 


6. Further dissection, if desired (at least segment VII 'should be 
removed) and final cleaning may now be carried out in preparation of clearing 
and mounting. 


STAINING 

Only a few general remarks regarding stains will be made here. 
Each worker has personal preferences and many stains are in popular 
use, among them add fuchsin, mercurochrome, and safranin. I have 
always found the first the most satisfactory. Add fuchsin is taken up in 
the greatest amounts by the sderotized parts of the integument leaving 
membranous parts dear. Thus sderites are clearly defined and easily 
observed and drawn. Tiny sclerotic processes in membranes also show 
up very distinctly. I have not found the other stains mentioned to act 
in this manner. 

Add fuchsin is an aqueous stain, i.e., it is soluble in water and must 
be applied to a fully hydrated specimen. I usually apply it after pre- 
liminary dissection using a 5% solution diluted 1 drop to 10‘ ml. 
Overnight is usually long enough to leave the parts in the stain. If not 
enough stain is taken up in this time more is added until the specimen 
is deeply tinted. Staining is always followed by destaining in distilled 
water to clear the membranes and obtain the desired differentiation. 

Non-aqueous stains may be applied to the specimens after they 
have been dehydrated in 95 % or absolute ethanol. 





FOR COLLECTING WITH LIGHTS 
12 1/2 LB POWER SOURCE 


WEIGHS 12-1/2 LBS. 

2 CYCLE, 1-3/4 HP 

EASY STARTING 

DIRECT DRIVE 

AIR COOLED 

MADE IN U.S.A. 

GUARANTEED 90 DAYS 

USES STANDARD OUTBOARD FUEL 

ROLLER AND BALL BEARINGS 

COMPLETELY SELF CONTAINED 

WORLDWIDE SERVICE AVAILABLE 

#5830 TINY TIGER GENERATOR 

115V AC, 12V DC, 300 WATTS $99.50 

#5831 CARRYING CASE 8.95 

Prices as listed are f.o.b. Santa Monica. 
California residents please add 4% sales tax. 




BIO METAL ASSOCIATES 

Box 61 

Santa Monica, California 


Volume 1 


Number 3 


March, 1963 


THE J0UIRNJAL ©F KESIA.^CH 
©NJ THE LEFIJ©©FTE^A\ 

IN THIS ISSUE 

Selection of allyl isothiocyanate by larvae of Pieris rapae and 

the inheritance of this trait 

William Hovanitz and Vincent C. S. Chang 169 

Biology of the Ceanothus stem-gall moth, Periploca ceanothiella 

(Cosens), with consideration of its control. J. Alex Munro 183 

Larval food-plant records for six western Papilios 

John F. Emmel and Thomas C. Emmel 191 

Colias philodice in Chiapas^ Mexico . . Thomas C. Emmel 194 

Notes on the early stages of two California geometrids 

John Adams Comstock 195 

Geographical distribution and variation of the genus Argynnis 

III. Argynnis diana William Hovanitz 201 

The generic, specific and lower category names of the nearctic 

butterflies. Part 2 — The Genus Colias . Paddy McHenry 209 

A standard method for mounting whole adult lepidoptera on 

slides utilizing polystyrene plastic . . Charles L. Hogue 223 


SPECIAL NOTICE 

The Journal is starting a series of short illustrated biographical sketches 
of past and present lepidopterists. In order to be wholly democratic and with 
no aim in mind of "honoring” anyone, all lepidopterists will be included. The 
only criterion will be a continued interest in the Lepidoptera, as shown by 
publication, collections, etc. The editor would appreciate receiving black and 
white prints of lepidopterists, together with a brief biographical sketch. This 
applies to lepidopterists of all countries. Authors sending manuscripts might 
send this material along with their articles. 



Volume 1 


Number 4 


Mny, 


THE JOURNAL 
OF RESEARCH 
ON THE LEFIJPORTERA 





a quarterly published at 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
edited by: WILLIAM HOVANITZ 

THE PURPOSE OF THE JOURNAL is to combine in one source 
the work in this field for the aid of students of this group of insects 
in a way not at present available. THE JOURNAL will attempt to 
publish primarily only critical and complete papers of an analytical 
nature, though there will be a limited section devoted to shorter papers 
and notes. QUALITY WORK on any aspects of research on the 
Lepidoptera is invited. Analytical and well illustrated works are pre- 
ferred, with a minimum of long description. 

AUTHORS ARE REQUESTED to refer to the journal as an 
example of the form to be used in preparing their manuscripts. Illu- 
strations should be of the best quality black and white, or line draw- 
ings and should be pre-arranged by the author to fit a reduced size 
of 4” X GVz." Footnotes should be avoided; bibliography should be as 
indicated. Tables should be set-up for page size as indicated. Manuscripts 
in good form and requiring little work by the editor will naturally 
appear first. Authors, who wish drawings made for them, may submit 
rough sketches and will be billed for the cost, which will be very 
negligible. Work to be done on research grants should so specify. When 
possible, tabular matter should be typed on good paper with a carbon 
ribbon in a form suitable for a one-third reduction and in a size to 
fit 4” X 61/2.” 

THE JOURNAL is not a suitable place for continued changes 
of nomenclature; unless the author is himself analytically studying a 
group from its biological point of view and finds a change necessary, 
the editor must ask authors to refrain from any changes from the 
McDunnough Check List unless superseded by a monograph published 
since that date. Popular books are not to be considered as giving scien- 
tific credence to any name. It is rare that name changes need be made 
and preference is given to old names unless in the editor’s opinion 
sufficient evidence is given to warrant such change. 

SUBSCRIPTIONS should be sent to the above address. 

RATES are: $8.00 per volume, personal subscription (but see below) 
$12.00 per volume, institutional subscription. 

The personal subscription rate is included in the membership to the 
Lepidoptera Foundation indicated below. SPECIAL SERVICE TO 
FOREIGN ADDRESSES: THE JOURNAL will be mailed air mail 
or registered at cost to the subscriber, if so desired. 


Journal of Research on the Lepidoptera 1 (4) ;237-244, 1963 

1140 IT. Orange Grove Ave,, Arcadia, California, U.S.A. 

© C&py fight 


TECHNIQUES IN THE STUDY OF 
POPULATION STRUCTURE IN 
PHILOTES SONORENSIS 

RUDOLPH H. T. MATTONI and MARVIN S. B. SEIGER 

Life Sciences Department, North American Aviation, Downey, Calif, and 
Department of Biological Sciences, Purdue University, Lafayette, Ind. 


The handling of living material has always been an im- 
portant factor in the design of biological experiments. This factor is 
especially critical in studying population structure under natural 
conditions. One of the most accurate techniques for determining 
parameters of population structure is that of capturing, marking, re- 
leasing and subsequent recapturing of individuals (Ford 1951). Such 
experiments would be biased if a behavioral response is elicited or 
differential viability imposed as a result of the technique employed. 

We have developed a handling technique in sampling populations 
of the small Lycaenid butterfly, Philotes sonorensis, over a period of 
three collecting years. This routine appears to have little or no effect 
on the subsequent behavior of the butterfly. The purpose of this 
report is to describe and evaluate the technique. 

Our objectives were to describe the distribution, numbers, and 
movements of adult individuals of P. sonorensis within a small circum- 
scribed area in the Fish Canyon portion of the San Gabriel Canyon 
Wash near Los Angeles. These individuals were classified as to 8 male 
and 5 female spot pattern phenotypes. (Figure 1). Six stations, each 
80 meters in diameter, were set up and sampled in 1955 and 1956. 
These were separated by distances ranging from 96 to 433 meters from 
the center of one area to the perimeter of another. The stations were 
destroyed by trenching operations in 1961 because of water require- 
ments. New experimental sites were established in 1963 in other 
areas of the wash. 

Sampling was done as weather permitted during the flight period 
in March. In 1955 a total of 809 specimens were captured 1126 times 
during 9 collecting days over a 21 day period. In 1956 there were 
972 specimens captured 1226 times for 11 collecting days over a 29 
day period. Sampling was done between 9 A.M. and 3 P.M., one 
hour being allotted to each station during each day. The order in which 
the stations were collected on consecutive days was ramdomized in 
order to minimize possible differential effects correlated with time 


237 


238 


PHILOTES SONORENSIS 


/. Ref. Lepid. 



-xi gJ Si 
£ 2 y ejOJS 
S 5-2 SH 
tJ ^-Q 2 

a <u >' 

JI3 O W5 

^ 2 > 

-ri *-< J3 ^ 

72 <u <u £ -s 

"O _Q “ U 

M " -H flj 

_Si iS S . n, 

= g cp g 

O 

Sv2 ^ « o 

a KH ^ 

“ §:sB 2| 

’w ^ ^ 

"g-2 o-c 

ia'S:3’|: 

o S< oj 2:5 I § 

« '-‘I Si bO c« 

> ° ^.2 s:: 

IjElll 

^ iZj . y rt> ^ 

13-c ^ 5 

c g-"^ o 
o m _ 


'ey a 


Os 45 

5^ o tu 

iS s 3 

s a 2 § 

C/D ' 1 -.H M_( WlH 


•- tyO 

-g G .S 
£ 


•.H ^ 

M U ‘ 

_Q cag 

Sy ' 

t 

IJJ 'S - 
C/D 52 f 


•13 a 

C5 Qj 
^ c/5 

V) -M 


•2 

§ 
O ' 
G . 
a> 
-G 
CU 


i(4):237-M4. 19^3 


AiATTONI AND SEIGER 


239 



Fig. 2. (below) Photograph showing equipment described in the text. 
RHTM on right holding a specimen about to be placed in the recovery 
chamber on the portable desk. Vials with specimens can be seen in the com- 
partment and the CO 2 tank behind. MSRS on left recording data. 

Fig, 3. (above) Anaesthesis Tube. 


240 


PHILOTES SONORENSIS 


/. Res. Lepid. 


of day. Each station was sampled for 30 minutes. This allowed 10 
minutes for moving between stations and 20 minutes for handling. 
When a butterfly was caught it was removed from the net with a 
T’ X 4 ” shell vial and the vial was plugged with a cotton stopper. We 
avoided touching the specimens with anything other than the net 
and the vial. After the collecting period the vials of butterflies were 
assembled in the center of the collecting area. In all cases one of us 
(RHTM) classified and marked the specimens while the other 
(MSBS) recorded the data. This process took approximately 20 minutes 
per station. The equipment used is shown in Figure 2. This included 
a portable desk, carbon dioxide tank, and release carton. The com- 
partment on the desk served to store vials in the shade. Each specimen 
was anesthetized in its vial by a 10 second exposure to CO 2 delivered 
at approximately 3 psi. The CO 2 was delivered from a small tank 
strapped to a pack frame for easy portability. A regulator maintained 
constant flow and pressure from the tank through the rubber tube 
to one of the glass tubes in a two hole rubber stopper inserted in 
the vial. The other glass tube in the stopper served as an exhaust 
to avoid excessive pressure in the vial ( Figure 3 ) • The anaesthetized 
butterfly was removed from the vial with flat bladed insect forceps and 
classified according to sex, forewing spot pattern and the area and 
date of previous capture if it had been a recaptured specimen. The 
specimen was then marked to indicate the date and area of capture. 
This was done by putting a dot of "Pactra” lacquer on the wing 
underside. The critical factor of this operation was maintaining a 
proper paint consistency. This was done by trial and error, using 
acetone as a diluent with a blunted dissecting needle. Six different 
colors were used to denote the six stations. These were applied to 
one of ten distinct underwing areas to denote the date of capture 
(Figure 4). The butterfly was then carefully laid on the bottom of 
a one gallon ice cream carton and allowed to recover. Recovery time 
varied, but seldom exceeded two minutes. The process was then 
repeated. After handling the last specimen, all the gear was assembled. 
Just prior to moving to the next station the carton was held upright, 
facing the sun, and was gently tapped so that the remaining specimens 
would fly off. If any specimen remained, the carton was inverted and 
vigorously tapped. If a specimen was not able to fly "normally” a 
distance of 10 feet, it was removed from the population and the 
event recorded. After the first day’s collection, subsequent collecting 
within about 10 feet of the center of the station was avoided as 
a precaution again recapturing injured specimens, if any should 
exist. In 1955, 21 individuals including 10 recaptures, and in 1956, 
37 individuals including 8 recaptures, were removed from the popula- 
tioa These figures are not wholly indicative of the effectiveness of 
the technique since about half of these represented specimens sampled 
for study. 


( 4 ) .- 237 - 244 . 19^3 


MATTONI AND SEIGER 


241 




Fig. 4. Marking scheme on underwings to denote data of capture. The 
circles indicate the positions of the lacquer marks on the wings which corres- 
pond to the collecting day. (designated by Roman numerals). 


Fig. 5. Recaptured specimens sampled on the last collecting day of 1955. 


242 


PHILOTES SONORENSIS 


/. Res. Lepid, 


The distinctive feature of our technique is the use of COq anaes- 
thesis in the field. We feel that anaesthesis greatly decreases the 
probability of damage due to handling, especially when examining 
and marking such a small butterfly as P. sonorensis. The procedure 
was adopted because of its general acceptability in the laboratory 
for research in insect physiology and genetics. For example, Seiger 
(1953) studied the effects of different anaesthetics on the dipteran, 
Drosophila mulleri. He found no significant difference in fertility and 
fecundity between flies which had been given no anaesthesis and 
flies which had been anaesthetized with COq. The average quiescent 
period after anaesthesis was one minute. This was sufficient time 
to classify an individual for sex and wing pattern, determine if it had 
been recaptured, mark the lower wings, and allow the lacquer to 
dry. Following recovery, the individuals tended to remain quietly 
in the container, thus further minimizing damage. The lacquer fastens 
the scales to the wing membrane and dries so quickly that no more 
than ten losses were suffered by insects sticking to the container or 
themselves. In the laboratory, P. sonorensis could withstand COq 
anaesthesis in excess of 10 minutes with no apparent ill effects, although 
under longer periods of exposure, partial paralysis would occur and 
eventually death ensue. There was no evidence for cumulative effect 
of repeated exposures of 10-30 seconds duration to COq. The possibility 
that abnormal behavior might result from anaesthesis has not been 
fully explored. In the first flight after anaesthesia, there appears to be 
a tendency for the butterflies to exhibit an escape behavior. After alight- 
ing once the behavior is not apparent. There appears to be no difference 
between the behavior of a butterfly in its first flight after anaesthesis 
and the behavior of a butterfly in flight after being captured in a 
net and released without anaesthesis. Although we feel that the advan- 
tages of anaesthesia far outweigh any possible disadvantages, we plan 
to determine whether there are any real effects of COq on the behavior 
of P. sonorensis in future experiments. 

We believe that the most important evidence of the negligible effect 
of our overall technique on behavior lies in the consistency of our 
data for two years with respect to the highly non-random pattern of 
movement. Another evidence was the remarkable behavior of 5 
individuals in 1956. These moved away from their area of capture 
and subsequently returned. There are several reasons which indicate 
that viability effects are also negligible. Figure 5 shows four marked 
specimens sampled on the last day of 1955. These appear to be quite 
undamaged, that at the upper right having been followed for eleven 
days and caught four times. In the course of our studies, several 
insects flew the distance between the two farthest stations (433 
meters), one flying a minimum of 819 meters. Lastly, because many 
specimens were captured more than once, it is possible to infer 
whether multiple handling had an additive effect on viability by com- 


i (4) is 37”244. 19^3 


MATTONI AND SEIGER 


243 


paring observed with expected values of multiple recaptures 0, 1, 2, 3 
and 4 times. 

Using a poisson distribution, the data for 1955 gives = 23.7 
and P = < .0001 for 3 df; and for 1956, = 5.4 and P = .12 

for 3 df. The highly significant departure from expected in 1955 
is due entirely to an excess of specimens recaptured 3 and 4 times. The 
meaning of this departure is obscure, although it does not contradict 
the hypothesis of additive deleterious handling effects. 

We found that the marking technique enabled us to distinguish 
individuals without confusion. The method can be compared to a 
punch card system in which each butterfly carries a recorded code on 
its wings, the color of the spot indicating the station where each 
capture took place and the position of the spot on he wing indicating 
the date of each- capture. Thus the population size for each station can 
be determined for any given collecting day (Dowdeswell, Fisher and 
Ford 1949 , Ford 1951), patterns of movement can be discerned and 
the life span of individuals can be calculated. These characteristics can 
be further quantified by correlation with maculation type and sex of 
the individuals. These data and conclusions are being prepared for 
publication in this journal 

For comparative information on field techniques in population 
study with Lepidoptera, the reader is referred to Abtot ( 1959), Ehrlich 
and Davidson (I960), Evans (1955), and Fales (1959); as well as 
those of Ford and his associates (op. dt.) We feel our technique 
has the least effect on viability and behavior differentials, particularly 
with reference to the use of anaesthesis; minimizing number of 
identifying marks on each individual; limiting individual contact to 
net, vial and forceps; and no more than 30 minute retention in the 
field. The last item, of course, is part of our experimental design. We 
feel that all such studies should take this factor into account, in spite 
of statistical difficulties in treating same day recaptures. 

We gratefully acknowledge the contributions of Mr. Jack Roper 
of North American Aviation in providing the photographs for Figures 
1, and 3, Dr. David Goodchild, CS & IRO for Figure 2, and Mr. 
Roy Pence, UCLA, for Figure 5. 

SUMMARY 

1. A method for anaesthetizing and marking individuals in order to 
determine population struaure is described. 

2. The benefits and possible disadvantages of the method are ex- 
amined and some applications of the technique are mentioned. 


244 


PHILOTES SONORmSIS 


J. Re$. Lepid. 


REFERENCES 

ABBOTT, W. 1959. Local Autecology and Behavior in Pieris protodice. Bois- 
duval and LeConte with some comparisons to Colias eurytheme Bois- 
duval, Wasmann ] Biol. 17: 279-298. 

DOWDESWELL, W. H., R. A. FISHER and E. B. FORD 1949. The 
Quantitative Study of Populations in the Lepidoptera. Heredity. 3: 67-84, 

ERHLICH, P. R. & S. E. DAVIDSON I960. Techniques for Capture — 
Recapture Studies of Lepidoptera populations. Lep. News. 14: 227-229. 

EVANS, W. H. 955. Retrieving marked Anthocharis reakertii. Lep. News. 
9: 118. 

FORD, E. B. 1951. The Experimental Study of Evolution. Australian and 
New Zealand Association for the Advancement of Science. 28; 143-154. 

FALES, J. H. 1959. A Field Study of the Flight Behavior of the Tiger 
Swallowtail Butterfly. Ann. Ent. Soc. Amer. 52: 486-487. 

SEIGER, M. B. 1953. The Effect of X-Ray Dosage on Fertility, Fecundity 
and Incidence of Chromosomal Aberrations in Drosophila mulleri. Master’s 
Thesis, University of Texas. 


Journal of Research on the Lepidoptera 1 (4) ;245-248, 1963 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 196 } 

EARLY STAGES OF A SOUTHERN CALIFORNIA 
GEOMETRID MOTH, DREPANULATRIX 
HULSTI HULSTI (DYAR) 

JOHN ADAMS COMSTOCK 

Del Mar, California 

Dr. Dyar, in 1904, named "Catopyrrha hulsti” from a specimen 
contained in the Hulst collection under another name, without desig- 
nating the locality of its origin. Dr. Rindge (1949) gave valuable 
data on the species, and its two subspecies, D. carneolata B. & McD. 
(1917) and D. verdiaria Grossbeck (1912) in his Revision of the 
Genus. He lists D. carneolata as an Arizona form of Drepanulatrix 
hulsti, and mentions California examples that seem indistinguishable 
from Arizona specimens. 

He places Drepanulatrix rindgearia Sperry (1948) as a synonym of 
D. hulsti. He also notes that the early stages are unknown. 

D. hulsti hulsti ranges throughout the greater portion of coastal 
California, wherever its food plant, Ceanothus, occurs. 

It flies most abundantly from May to August, but has been reported 
as early as January, and late into September. 

Eggs of this species were obtained this year (1962) in Del Mar, 
on several occasions, from May 25 to late August. For the most part 
these were laid singly on their sides. When first laid they were light 
green, with a pearly luster. Shortly they changed to cream or light tan, 
with contrasting dots and irregular tortuous bands of dark pink to 
salmon-red. 

The characteristic from is ovoid, with a flattened base and rounded 
tip, but some approach a barrel-shaped form with flattened base. Our 
illustration (figure 1 A to C) shows the extremes of this variation, 
and also the early and late changes in markings. 

Sculpture of the surface consists of longitudinal ridges, from 28 to 
36 in number, running from the edge of the flattened base to the top. 
These ridges have pearled’ edges, the pearls or raised points correspond- 
ing to the numerous horizontal lines crossing between the ridges. The 
raised pearls are small and relatively inconspicuous. 

On the flattened base the ridges end abruptly, and within the circle 
formed by their terminations the surface is covered by numerous 
irregular hexagonal pits with raised walls, as shown in figure IB. 

There is considerable variation in the size of eggs, the average being 
1 mm. tall by 0.6 mm. wide. Eggs laid May 25 hatched June 3. The 
larval egress was through one end, the remainder of the shell being 
left intact. 


245 


246 


JOHN ADAMS COMSTOCK 


/. Res. tepid. 


By comparison with the egg of Drepanulatrix monicaria it is larger, 
the number of ridges greater, and the salmon-red markings heavier. 

FIRST INSTAR LARVA: Length, 375 mm. Head width, 0.4 mm. 
The head is wider than the first segment. The ground color is white 
with a heavy spotting of black dots on a brown base each side of the 
epicranial suture and the outer and upper two-thirds of each cheek. 
There are two raised papillae on the front. The mouth parts are brown, 
and the ocelli black on a white ground. 

The first thoracic segment is narrower than the head, but wider 
than the remaining segments. The ground color of all body segments 
is white. There is a longitudinal middorsal irregular band of olive which 
is somewhat restricted at the segmental junctures. A narrow irregular 
dorso-lateral band parallels it. There are several rows of black papillae 
running generally in a longitudinal direction. The placement of these 
on the dorsal surface is shown in the illustration, (figure ID). Each 
papillus is surrounded at its base by a white circlet, and mounts a 
black seta at its tip. 

The legs are hyaline, with a tinge of light yellow. The single pair 
of prolegs and the anal prolegs are concolorous with the body. 

LARVA OF 15 MM. LENGTH: Head width, 1.5 mm. Head, gray, 
spotted with black, the spotting heaviest on the margin of the crown 
and cheeks. The front is slightly less spotted. The mandibles and ocelli 
are black. 

The body ground color is gray. Most of the segments are crossed 
transversely by five or six folds, on which are placed prominent black 
papillae bearing black setae. The entire surface of the body is heavily 
sprinkled with minute black and brown dots. These have a slight ten- 
dency to form longitudinal lines. The spots on the ventral surface are 
predominantly brown and are more definitely arranged in longitudinal 
lines. 

The prolegs are concolorous with the body, and the true legs are 
less spotted and more hyaline. 

MATURE LARVA: Length, 26 mm. Head width, 2.2 mm. Head; 
color, gray-brown, heavily spotted with minute brown dots. The front 
is finely ridged horizontally, with a line of six minute black dots running 
transversely near the lower edge. The labrum is darker, and is ridged 
longitudinally. The maxillae are dark brown, the ocelli black, and the 
antennae translucent. 

Across the center of each cheek is a lunate black band, beginning 
near the epicranial suture, extending laterally, then arching inferiorly 
to end near the ocelli. The setae are brown, and arise from black papillae. 
The head is shown in front view on figure IE. 

BODY: Ground color, whitish gray, nearly obscured by numerous 
small brown and black dots. There is a suggestion of a double middorsal 
longitudinal brown stripe in the thoracic area. A poorly defined dark 
spiracular band, made up of dots, is present on some specimens. The 
spiracles have narrow black rims and cream colored centers. The legs 


( 4 ) : 245-248 


JOHN ADAMS COMSTOCK 


247 



Figure 1 

Early stages of Drepanulatrix hulsti hulsti. A. Egg, lateral aspect final 
color phase, enlarged X 40. B. Base of egg, enlarged X 40. C. Egg. elongate 
form, and early light phase, enlarged X 40. D. First instar larva, dorsal aspect, 
enlarged X 21. E. Head of mature larva, front view, enlarged X 10. F. and 
G. Mature larva, dorsal aspect, enlarged X 5.5. H. Pupa, ventral aspect, en- 
larged X 6. I and J. Cremaster, ventral and lateral aspects, enlarged X 14. 

Reproduced from water color drawing by the author. 


248 


GEOMETRID M.OTH 


/. Res. Lcpid, 


and prolegs are translucent, the latter being spotted with brown. The 
crochets are red-brown, apparently biordinal in alignment, and developed 
only along the lateral edge of the oval foot-pad, with short stubs only 
on the medial edge. The setae are light brown, relatively short, and arise 
from minute black papillae. ( figure 1 F and 1 G ) . 

Pupation occurred on the floor of the rearing jar, among leaves and 
debris, mixed with a few strands of silk. 

PUPA: Length, 14 mm. Greatest width, through middle of wing 
cases, 4.2 mm. 

The anterior two-thirds of the pupa is relatively wide and plump. 
The six terminal segments taper sharply to a distinctive cremaster, as 
will be noted on figure 1 H. An enlargement of the latter is pictured on 
figure 1 I. 

The eyes are prominent, dark, and protruding. The cephalic end 
is evenly rounded. The antennae and maxillae extend to the margins of 
the wing cases. The spiracles are relatively small and nearly indisting- 
uishable without a lens. There are no setae. 

The color of the pupa is predominantly reddish brown, but the eyes 
and cremaster are black. 

The cremaster is pear-shaped, with the stem end narrowed and 
elongated. Laterally it bears two globular bodies, one on each side, when 
viewed in ventral aspect. Portions of the surface are nodular. The tip 
bears several short spicules which arch dorsally. Two of these are 
longer, and recurve ventrally. These details can best be grasped by 
referring to the illustration on figure IH. 

The texture of the body surface of the pupa is predominantly smooth 
and glistening. 


BIBLIOGRAPHY 

BARNES, WILLIAM and J. McDUNNOUGH.,1917. Contributions to the 
Lepidoptera. 3 (4) : PI. 28, Figs. 5-6. 

DYAR, HARRISON G. 1904. A few notes on the Hulst collection. ?roc. 
Ent. Soc. Wash. 6: 226. 

GROSSBECK, J. A. 1912. Jour. N. Y. Ent. Soc. 20: 285. 

RINDGE, FREDERICK H. 1949. A revision of the Geometrid moths 
formerly assigned to Drepanulatrix. Bull. Am. Mus. Nat. Hist. 94 (5) : 
233-298. 

SPERRY, JOHN L. 1948. Three apparently undescribed Geometrid moths 
from the southwest. Bull. So. Calif. Acad. Sci. Al (1): 6-7. 


Journal of Research on the Lepidoptera 1 (4) :249-259, 1963 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A, 

© Copyright 196} 


THE EFFECTIVENESS OF DIFFERENT ISOTHIOCYAN- 
ATES ON ATTRACTING LARVAE OF PIERIS RAPAE 

WILLIAM HOVANITZ; VINCENT C. S. CHANG AND 
GERALD HONCH 

The California Arhoretum Foundation and the Lepidoptera Foundation, 
Arcadia, California; The Los Angeles State College; and 
The University of California, Riverside, California 


The effectiveness of black mustard oil (allyl-isothiocyanate) 
on attracting larvae of Pieris rapae at various concentrations has already 
been described in this JOURNAL (Hovanitz and Chang, 1963). 
This mustard oil is the commonly used oil for condiments and is pre- 
pared by pressing from the seeds of black mustard {Bras ska nigra). 
This oil is known also from various species of Cruciferous plants in 
addition to black mustard. Other mustard oils are known from a wide 
variety of plants, not all of them members of the family Cruciferae. 
This paper describes some experiments on the selection by Pieris rapae 
larvae on allyl-isothiocyanate produced by two different methods, and 
on three other isothiocyanates. 

NATURALLY OCCURRING ISOTHIOCYANATES 

Isothiocyanates (mustard oils) are naturally derived from gluco- 
sides which occur in a wide variety of the higher plants, belonging to 
a relatively small number of plant families (Kjaer, I960). These 
glucosides are characterized by the ability to undergo enzymatic hydro- 
lysis to isothiocyanates, hyrogen sulfate and D-glucose. The latter 
has invariably been encountered as the sugar moiety of the more than 
thirty individual compounds recorded thus far (Kjaer, I960). Accord- 
ing to this author, only nine glucosides of these thirty isothiocyanates 
have as yet been crystallized. The occurrence of more than one gluco- 
side in a given plant species is most common; as many as eight in- 
dividual glucosides have been distinguished in a single seed specimen. 
The compounds appear to be distributed over the entire plant; the 
glucosides are diffusely present in parenchymal tissues, especially in 

^ Aided by a grant from the National Science Foundation, Washington, D. C. 


249 


250 


HOVANITZ, CHANG AND HONCH 


/. Res. Lepid. 


the bark (Guignard, 1890, 1893). The embryo constitutes the site of 
accumulation in seeds. Little has been done to relate the variation in 
glucoside content as a function of the stage of growth, or of environ- 
mental factors such as climate, soil, etc, Stahmann, et al (1943) found 
that 2-phehylethylisothiocyanate, the aglucone of gluconosturtiin, to be 
the predominant mustard oil enzymatically liberated from the roots of 
Brassica nigra, whereas the seeds of this plant represent the classical 
source of the glucoside sinigrin which yields allyl isothiocyanate upon 
enzymatic fission. Delaveau (1958) has noticed considerable varia- 
tion in the total and relative amounts of the individual glucosides in 
the plant Alliaria officinalis during its growth cycle. Considerable vari- 
ation has been detected in quantitative and qualitative differences in 
glucoside content of various parts of a plant as well as in different 
lots of the same plant tissue. 

A list of the known natural isothiocyanates and their parent gluco- 
sides in various plant genera and species which are known to be hosts 
of species of Pieris is given in table 1. In this table the plants are listed 
by families and genera. After each genus or species of plants is listed 
( 1 ) the Pieris known to feed on it, ( 2 ) the mustard oil glucoside 
which has been isolated, or established as having been found, in that 
plant, (3) the aglycones (the mustard oils corresponding to the gluco- 
side) and (4) the group of compounds to which it belongs. 

Observation of this list does not seem to indicate any common 
denominator between the mustard oil, or glucoside, and the species of 
Pieris which is attracted to it, with a single exception. That is, sinigrin 
is present in most plants attracted by one or more species of Pieris. 
This, in itself, may mean little for sinigrin appears to be the most com- 
mon mustard oil glucoside, and the earliest known. There are Pieris 
attracted by plants in which sinigrin is not known, for example, 
( 1 ) Lepidium, which is a common food plant of Pieris protodice and 
P. occidentalis, (2) Cleome (and probably Isomeris), which is a com- 
mon food plant of P. protodice, and P, beckeri, ( 3 ) Propaeolum, a food 
plant of Pieris rapae, and (4) various Resedaceae, food plants of Pieris 
daplidice. In each of these plants indicated, there are some other mus- 
tard oil glucosides present, namely, glucotropaeolin and glucoraphanin 
for Lepidium, glucotropaeolin for Tropaeolum, gluconasturiin, gluco- 
barbarian, and glucotropaeolin for the Resedaceae, and glucocapparin 
for Cleome (or Isomeris, Capparidaceae) . 

It seems likely that more than one mustard oil attracts Pieris, 
even of the same species. The possibility exists that there may be little 
specificity for the kind of mustard oil, so long as some one isothio- 
cyanate is present in the proper concentration. Except for Tropaeolum, 
all listed food plants of Pieris rapae contain sinigrin in some part of 
the plant. Only glucotropaeolin is known from Tropaeolum itself, yet 
Pieris rapae survives on that plant after only a few generations of adap- 
tation. 


i (4):249-259, 1963 


ISOTHIOCYANATES 


251 


TESTS OF LARVAE OF PIERIS RAPAE TOWARD 
VARIOUS ISOTHIOCYANATES 

The tests to be described here were made by subjecting larvae of 
Fieris rapae from our laboratory strain, grown for several generations 
on black mustard {Brassica nigra), to our testing device. This device 
has been described previously (Hovanitz and Chang, 1963). It con- 
sists essentially of a greenhouse flat, with filter paper moistened with 
dilutions of mustard oils, and spaced equally around the periphery of 
the flat. The larvae are set in the center and make their way to the 
side of the flat, being attracted to one or another of the mustard oils, 
at various concentrations. 

For this experiment, tests were made of five different mustard oils 
in concentrations from 10~^ to 10~'^. These five were: 

( 1 ) benzyl-isothiocyanate : this is the aglycone of glucotropaeolin, 
found in Cruciferae: Lepidium sp., Resedaceae, and Tropaeo- 
laceae; Tropaeolum sp. Synthetic origin. 

(2) phenyl-isothiocyanate: there is no natural aglvcone of this 
mustard oil listed by Kjaer. Synthetic origin. 

(3) phenethyl-isothiocyanate: this is the aglycone of gluconasturti- 
in, found in Barharea, Brassica nigra. Brassica oleracea and 
the Resedaceae. Synthetic origin. 

(4) allyl-isothiocyanate: this is the algycone of sinigrin found in 
many plants. Commercial synthetic. 

(5) same as above, only commercial natural product from Bras- 
sica nigra seeds. 

Sources of the above mentioned mustard oils are described below. 

General Procedure for the Preparation of Synthetic Mustard Oils 

The following general procedure for the preparation of mustard oils from 
various amines was provided by Professor Henry Klostergaard of the San 
Fernando Valley State College. This method was used for the preparation of 
the phenethyl-, benzyl-, and methyl-isothiocyanates used in this paner. All 
procedures to be carried out at 0°C. 

In a 2000 ml. Erlynmeyer flask, mix 0.07878 moles of the amine and 
enough absolute ethyl alcohol to make a 30% solution ( by volume). To 
the above solution add 4.8 ml. CS^ solution (0.03939 moles in a 1:1 solution 
with absolute alcohol). Wait 10 to 15 minutes, stirring often as there will 
be some heat evolved. Now add 100 ml. of the iodine solution (0.03939 
moles or 10 gms. of iodine dissolved in 100 ml. absolute alcohol). The 
solution will start to clear up. Disregard any crystalliztion at this point. 
Shake until the brown color is gone and some yellow color appears. This is 
not stable so immediately go on to the next step. At this point add 25 ml. 
of the sodium solution (0.03939 moles or 0.9 gms. of sodium in 25 ml. 
absolute alcohol). The mixture will split and become homogeneous. 

Now add another 100 ml. of the iodine solution as above. Some ele- 
mentary sulfur will settle out (this is not soluble in ether). Now add 250 ml. 
acidifed water (1% HCL). 

Next pour the solution into a separatory funnel and extract the mustard 
oil with ether. The mustard oil will appear in the ether phase or the top 


TABLE 1. The occurrence of natural isothiocyanates and their parent glucosides in plant 
and species known to be hosts of species of Pieris (chemical data from Kjaer) . 


252 


HOVANITZ, CHANG AND HONCH 


J. Res. Lepid. 







(U 

a 

u 





glucorapiferum 2-Hydroxy-3-butenyl 

isothiocyanate 


19^3 


ISOTHIOCYANATES 


253 



•S 8 g g g § 
’£3 o 3 


> 3 

© 0) G) EQ 

"8 © © *^ 

^ ^ ^ 
gj ^ ^ 4^ 

0) OJ 

^ 0) 0) S 


© © © ^ © 






Q a 

© ^ u U u 
•H ^ -H O ^ -H 
^ O.^ -H d ^ 




m 

d S 

5^ 


§ g 


§“^ 


M U m O 

4J a d ® 

© © i 

m 65 ^ i 
=H «H O 

>, S'ja 3 
& o ■■ "^ 


« 'S 


U I 

d 


© >» CO © 

§ g”" § 
"S'© 'S 
0 ^ 0 


I 3 


d £0 a *3 

W ^ »H M 

^ ^ a d 

§5 S^SrS 

O © O 60 O 

O O © »H U 

d d 3 d d 

60 60 60 sa 60 


3 d 

pH »H 
60 m 


§ 

'S. d 


3 d 
60 m 


•H I. d 1^ 

»S^ CO -f>! eg 
d «H M ^ 
® CO ^ fH 


3 d 

fH 's=i 
60 05 


d 

•H 

1-5 M 
3 cd 
4J ^ 
CO U 

ed d 
d ^ 



Tropaeolum P . rapae glucotropaeolin benzyl isothiocyanate aromatic isothiocyanates 


254 


HOVANITZ, CHANG AND HONCH 


/. Res. Lepid. 


layer. Discard the lower layer. Add some 10% NaOH to the mustard oil 
and ether solution to take off the iodine and then discard the lower layer. 
Next wash the solution with an equal volume of distilled water. Discard 
the water layer. Now there is present an ether solution of the mustard oil, 
(Ice this solution immedately.) Vacuum distil this solution in a flash evap- 
orator with the solution in the flask cooled by an ice-salt solution and suc- 
cessive flasks cooled by ether and dry ice. Keep the mustard oil and ether 
solution under vacuum for at least IVz hours. The mustard oil will be left 
in the original flask. Remove it immediately and store at 0°C. 

Caution 

All the solutions involved in this preparation should be kept refrigerated 
at 0°C. until used. All work should be done under an adequate hood since 
the chemicals involved are highly toxic and odoriferous. 

Tests of the larvae of Pieris rapae as indicated were made separately, 
utilizing the five dilutions of the mustard oils indicated. The actual 
dilutions may actually have been less than indicated because it appears 
that at the temperature used (about 20 °C.), not all the mustard oil 
may have gone into solution. Each larva was tested twenty times, giving 
total test times ranging from 180 times for benzl-isothiocyanate to 
680 times for phenyl-isothiocyanate. 

The allyl-isothiocyanates differed in their response, according to 
their origin. One was derived from black mustard seeds by standard 
commercial techniques of compression, and fermentation. The other 
was synthetically prepared. In each case, the larvae selected the 10“*^ 
dilution of mustard oil (22.1%, for synthetic to 21.2% for natural) 
but the dispersal of the selections was different. For example, the 
dilution of 10~^ was selected by the larvae for the synthetic mustard 
oil 20.7% of the time, as compared with only 11.6% for the natural 
product. Dilutions of less than 10~^ are also different. For example, 
while the synthetic gave selections of 193% at 10~®, the natural 
product gave 14%. At 10~'^, these were 15.2% for the synthetic as 
compared with 11.6% for the natural. An apparently significantly 
greater number of larvae left the flat without any selection at all with 
the natural (24.2%) as compared with the synthetic product (2.5%). 
This discrepancy was probably due to the human variation in carrying 
out the tests. The percentage is greater when the larvae are not allowed 
as long a selection period. This discrepancy would not affect the actual 
selections themselves. 

Another effect is quite noticeable with these data, as well as with 
the data on the other mustard oils, namely, that the selections give a 
bimodal curve (see fig. 1). Such a bimodal curve was not clear with 
our previous testing though a trace of it might be detected (fig. 3, 
Hovanitz and Chang, 1963). The previous testing was made with 
the natural product which also shows a very poor bimodal curve ( fig. 1 ) . 
The synthetic product shows a more strongly indicated bimodality at 
10“'^ and at 10~® (fig. 1). The reasons for the bimodal curve which 
is especially apparent for the synthetic allyl mustard oil are unknown. 

Selections for benzyl-isothicoyanate are almost certainly negative. 


TABLE 2. The selection by larvae of Pieris rapae for various concentrations 
of mustard oils. 


(4):249-259, 1963 


ISOTHIOCYANATES 


255 


CO 






T— 1 rH 






cC cd 

0 

0 

0 

0 

0 

4J -H 

00 

CO 

0 

NO 

0 

0 J-) 

rH 

NO 

<[■ 

m 

m 

U 






M-! 0) 






0 c8 






> 






• }-i 

ON 


0 

00 

m 

0 cS 


cn 

CM 

CM 

CM 

C <-* 








6^ 



B^ 

C 


<1- 


in 

CM 

0 

gvO 


cn 

• 


•H 

LO 

cn 


CM 

<t 

4J 


rH 

nD 


CM 

0 






0) 






0 r-H 

rH 

tH 

m 



fl 0) 

00 

ON 

CM 

rH 

CM 

CO 





I— 1 





B^ 






CM 


T3 

6^ 

CNJ 

B^ 


B^ 

0) 

<!■ 

• 

00 

0 

NO 

rH $-1 


CM 


pH 


r-l QJ 

ON 

pH 

NO 


NO 

•H 4J 






4-t cd 






M & 


tn 



CO 

•H 

rH 

CO 

CM 

LO 

CO 








6-S 


B-5 

5^ 

B^ 


00 

a^ 

00 

CM 

vD 


cn 

?H 

00 

m 

rH 

ON 

rH 

r-l 

1 — 1 

iH 

tH 

0 






iH 

U-l 

rH 

m 

LO 

00 


CS) 

00 

NO 

00 

LO 





B^ 






CO 



CO 








as 

ON 

pH 

00 

CO 

rH 

rH 

f — 1 


0 







in 

CO 

vD 

00 

0 



rH 


0 










5^ 

B^ 

B^ 

B^ 



00 

m 

0 

00 


fH 






• 

T— 1 

CO 

0 

0 



I—l 

i—i 

r— 1 

r-4 

*0 






rH 

sH 

0 

00 

NO 



sH 

00 


MO 

m 



B-S 



B^ 




B^r 

, — [ 

CM 


CO 


00 





00 


CM 

1 — 1 

NO 

00 

»H 

0 

CM 

CM 

1 



CM 



0 







m 



<1- 

NO 


rH 

CH 

CO 

CM 

0 



rH 

00 

I — 1 





B^ 

6 ^ 




6^ 

0 


v£> 


ON 

<5- 







m 

0 

pH 


00 


I — I 

CM 

iH 

u -1 






0 

nD 

r-H 

0 

vD 

00 

r-i 

t-H 

0 

NO 

iH 

m 





rH 











1 

0 





r-H 





1 

>N 1 

1 4J 

1 


1 0 0) 

1 0 cu 

0 0) 

0 0) OJ 

0 d rH 


^ 

t-M -H 4-1 

4-1 -H 4J 

1 -H 4 J 43 

1 -H 4-1 Cd 


cd 

>N,r: Cd 

0) 43 Cd 

rM 43 Cd 4-J 

rH 43 cd d 


N 4-> d 

d -u d 

P 3 4 J d 

>N -u d c 3 

>N 4J d 3 


d 0 cd 

(U 0 cd 

0) 0 cd 

rH 0 cd >, 

1-4 0 Cd 4-1 


0 ) CO >, 

CO 

43 CO 

r-H CO W 

rH CO tn Cd 


PQ -H 0 

PM -H 0 

CM -H 0 

■< -i-l 0 

<d “H 0 d 


PERCENT PERCENT 


256 


HOVANITZ, CHANG AND HONCH 


/. Res. Lepid. 


25 




Fig. 1. Histograms illustrating the fomparison between natural and snythetic mustard 
oils respecting their selection by larvae of Pieris rapae. Note the strong bimodality of the his- 
togram showing the synthetic product as compared with the natural 


PERCEN T PERCEN T PERCEN T 


(4);249-259, 1963 


ISOTHIOCYANATES 


257 


- 

— 


BENZ) 

YL 





10 - ® 


10 ^ 

10 '® 

10-3 

HgO 

NONE 


f 




Fig. 2. Histograms illustrating the selections made by larvae of Picris rapae for various 
dilutions of the mustard oils benzyl-isothiocyanate, phenyl-isothiocyanate and phenethyl-ioso- 
thiocyanate. Note the bimodality of two of the histograms and the nearly complete lack of 
selection indicated by the other. 


258 


HOVANITZ, CHANG AND HONCH 


/. Res. Lepid, 


not varying significantly from water alone (table 2 and fig. 2). This 
mustard oil also was the one in which dissolving in water appeared 
almost impossible and in which case the tests might very well have 
been conducted on water alone. Perhaps at a higher temperature, dis- 
solving the mustard oil might have been accomplished but under the 
circumstances it does not seem as if this mustard oil could be of any 
influence on attracting Pieris rapae larvae to Tropaeolum species where 
it is found naturally. 

Phenyl-isothiocyanate and phenethyl-isothiocyanate were about 
equally effective in attracting the larvae though there is a possibility 
that the phenyl mustard oil may be slightly less attractive than the 
phenethyl, since at each concentration the percent attraction was 
slightly less. However, the decrease possibly is not significant. The 
attraction of both of these mustard oils was not much different from 
that of allyl isothiocyanate (table 2 and fig, 2). This leads to the 
interesting possibility of the specificity between isothiocyanates in the 
generic sense and the Pieridae that are attracted to plants that contain 
the compounds. It is possible that the attraction is to the isothio- 
cyanate part of the molecule and not to side chains. "When applied 
to the tongue, all mustard oils cause a sharp and burning sensation. 
Their odors, though mostly pungent, display characteristic individual 
differences which often are helpful in the detection and classification 
of mustard oils. Certain isothiocyanates, that undergo rapid intramole- 
cular cyclization, give rise to a transient biting taste, followed by a 
sensation of bitterness. Like most synthetic mustard oils, those of 
natural origin show vesicant and frequently also lachrymatory proper- 
ties,” from Kjaer (I960). These same substances, however, when 
present in small concentration appear to give rise to the desirable con- 
diment properties of the mustard oils and appear in turn account for 
the specificity of these oils to the larvae and adults of Pieris. 

CONCLUSIONS AND SUMMARY 

1. A list is given of the mustard oils which are known to occur in 
food plants of the Pieridae. These are derived from naturally-occurring 
glucosides which are also listed. 

2. It seems evident from this list that more than one mustard oil 
attracts Pieris, since the evidence does not indicate that only one could 
be responsible. 

3. Tests were made on the selection of larvae of Pieris rapae 
toward several mustard oils at various concentrations. These oils were 
either commercially obtained, or were synthetically compounded by us. 
The procedure used for synthetic compounding is described. 

4. Synthetic and natural allyl mustard oils obtained commer- 
cially gave somewhat divergent testing results. The synthetic gave a 
bimodal curve with modes at 10”^ and 10”® which were only slightly 


( 4 ) 5249 - 259 , 19^3 


ISOTHIOCYANATES 


259 


or flot at ail apparent with the natural. 

5. Other mustard oils tested were benzyl-isothiocyaoate, phenyl- 
isothiocyanate and phenethyl-isothiocyanate. 

6. Benzyl-isothicoyanate is the aglycone of glucotropaeolin, the 
glucoside of Tropaeolum sp. and some other plants. Its attraction power 
at the concentrations used is not different from that of distilled water. 
In view of the difficulties experienced in dissolving this substance^ it 
is possible that none actually dissolved and that, therefore, the tests were 
actually on water alone. 

7. Phenyl-isothiocyanate is not known to be natural occurring. 
Nevertheless, it had a selective effect on the larvae only slightly less 
effective than allyl- and phenethyl-isothiocyanates. 

8. Phenethyl-isothiocyanate had an attractive power almost equiv- 
alent to that of the synthetic allyl-isothiocyanate. It is a naturally- 
occurring mustard oil, being the aglycone of gluconasturtiin which is 
found in Nasturtium and Barbarea. 

9. It appears that more mustard oils than just allyl-isothiocyanate 
are attractive to Pieris. The experience with benzyl-isothiocyanate 
however indicates that not all are effective, at least not under the ex- 
perimental conditions utilized. 

LITERATURE CITED 

DELAVEAU, P. 1958. Variations de la teneur en heterosides a senevol de 
VAUiaria officinalis L. an corns de la vegetation. C. R. hebd. Seances 
Acad. ScL 246: 1903. 

GUIGNARD, L. 1890. Recherches sur la localisation des principes aaifs des 
Cruciferes. /. Botanique 4: 385. 

1893. Recherches sur la localisation des principes actifs chez les 
Capparidees, Tropeolees, Limnanthees, Resedacees. /. Botanique 7: 345. 
HOVANITZ, W. and V. C. S. CHANG. 1963. Selection of allyl-isothiocyanate 
by larvae of Pieris rapae and the inheritance of this trait. /. Res. Lepid. 
1(3) : 169-182. 

KJAER, ANDERS. I960. Naturally derived isothiocyanates (mustard oils) 
and their parent glucosides. Fortschritte d. Chem. org. Naturst. 18: 
122-176. Springer, Wien. 

STAHMANN, M. A., K, P. LINK and P. C. WALKER, 1943. Mustard oils 
in crucifers and their relation to resistance to clubroot. /. Agric. Res. 

67: 49. 











N 

k- 

rt ■ '*, ' 

Ul? ^■'3^'■^■.1tW‘■1 C^T; ir*’-" ' w:dihj.v -;■■*>'! *V I '4^l)55 -'-V *’ V 



1 




vi! /i r :,'V-' .-^,. 

I .?■ " ‘^KtfTd/, !* Stoi- 
T* ^ ! k:c.. jal*'< ‘ ■ 

,■’ f _ '' . ll;ni*iirjr!'ai *i 




tat,... 


’. 4- ,t?4»^5^‘>i^.^', ^r‘.w‘*-..'^V; 
t .■*'»’ » ’*• rlt * vrtTt^ ■•> ti ■ w 

•;.^^■'^^^y•4'i^>^■ *j-VsVll 
' " ■ ■ * -'^.V»,.'^ 5. ' . -C:. 

■• vj- ■!'’• 

u<^‘ %:!', ••■■;)' i ... 

• ' . ■ ■■ *.■: ■■ _ „ 

■ ' ■ *t irtfi* yt ' ■ 

^ ■ >/ .. - ^ 

V', .V*rjg3*'W*»vi“ 




TA 









. 

JtT^' yJf.^(^^^'SW^ffT'’ WI-%V<‘' -.’tr’i ..t-H'..' .r '■ •“'rr- 

a .s... -fr it^fMr;ri->.-; . ^M'.‘fi-- .-.I.v - . ■§ 









" . ' r^t ... [ Cl i 



Journal of Research on the Lepidoptera 


1(4) :261-274, 1963 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
© Copyright 1 ^ 6 } 


THE ORIGIN OF A SYMPATRIC SPECIES IN COLIAS 
THROUGH THE AID OF NATURAL HYBRIDIZATION 

WILLIAM HOVANITZ 

The Lepidoptera Foundation and the California Arboretum 
Foundation, Inc., Arcadia, California and the Los Angeles 
State College, Los Angeles, California 

1. INTRODUCTION 

The theoretical set up for the origin of a new species in any 
biological group is the existence of two or more species in the same 
geographical area, which mix and reassort their combined gene com- 
binations by the process of hybridization, Fi intercrossing and back- 
crossing. There have been examples described in recent years in which 
crossing of this sort has served to blend together two existing species 
in a given area in such a way that both parental species cease existence 
as independent entities; this situation has been described generally as 
"subspecific intergradation” because of necessity the blending must be 
done on a geographical, or spatial, scale. Recent examples of this have 
been described by Sette (1962) involving the subspecies macaria and 
laufina of Argynnis callippe, and others by Hovanitz (1949). 

In those cases in which natural hybridization of the two species has 
served only to permit gene (and character) interchange between the 
two original species, both of which maintain themselves as independent 
entities, but in which no new species is formed, the situation is described 
as one of introgressive hybridization. One of the most extensive cases 
of introgressive hybridization known is that existing between the 
species Colias emytheme and Colias philodice (Hovanitz, 1943, 1944a, 
1944b and 1948). Other cases which are less extensive are known in 
many other species of Colias. 

Much more rare, possibly because the occurrence may be transient, 
is the situation in which two hybridizing species retain their separate 
existence, but out of the assorted gene pool arising by hybridization, 
there come a third species which maintains itself as a separate entity 
without the further necessity of hybridization. This is the situation to 
be analyzed in this paper. 

It is almost axiomatic that for closely related and interfertile species 
to exist in the same general area, and yet to maintain their separate 
identities, those two or more species must be separated in some way 


261 


262 


HOVANITZ 


/. Res. Lepid. 



Fig. 1. North polar projeaion map showing the world distribution of 
Colias hecla. 

hy physiological or ecological isolation barriers for if this were not so, 
the genes of the two or more species would be blended together as in 
one gigantic melting pot. The isolation barriers for Colias appear to be 
such physiological and ecological ones rather than genetic ones since 
most species appear to be interfertile. Tied with these barriers is the 
selection for genes controlling adaptation for different habitats and dif- 
ferent food plants of the larvae, which enable Colias to exist under a 
greater variety of conditions than would be possible were they all 
amalgamated into one species in a given geographical area. 

Previous study has indicated that there is considerable natural 
hybridization in the genus Colias (Hovanitz, 1949, 1956). The species 
Colias philodice and Colias eurytheme have been shown to hybridize 
rather freely throughout all parts of their ranges where they are in 


i(4):i6i-274. 5963 


SYMPATRIC SPECIES 


263 



\ ■ ■ —I-- , i ■■ ■ »■ ' ^ Colios nQSt0S l-AWBERT'S AZIMUTHAL EQUAL-AREA PROJECTION 

Colias phicomone 

Fig. 2. North polar projection map showing the world distribution of 
Colias nastes. 


contact, or where there is overlap of their habitats (Hovanitz, 1944a, 
1944b). This is a classical example of intro gression, the reciprocal 
transfer of genes between two or more species in natural contact, but 
without the complete submergence of the identity of any. Similar 
examples of natural hybridization have been described for the following 
species combinations of Colias: 

Colias interior - C. christina 
Colias hyale - C. erate - C. croceus 
Colias Christina - C. gigantea 
Colias philodice = C. eurytheme 
Colias hecla - C. nastes 
Colias interior - C. philodice 
Colias alexandra - C. philodice 
Colias Christina - C. philodice 


264 


HOVANITZ 


/. Res. Lepid. 



Fig. 3. North polar projection map showing the world distribution of 
Colias palaeno. 


Without doubt a great number of additional combinations will be 
found, wherever any two or more of the species of this genus occur 
together. Species hybridization in the laboratory, or in the field, lead- 
nig only to Fj generation individuals, with no history of fertile back- 
crossing or of fertile Fo production, would have no effect on altering 
the composition of natural populations. Such crosses therefore are 
only indicative of physical ability to mate, of Fi to survive (though 
not of Fi fertility), and to indicate something of the nature of genetic 
dominance of the many characters involved in the cross; however, they 
give no information at all on the evolutionary relationships between 
the parental species. Further testing must be done for this purpose by 
carrying the crosses through to the F2 generation and backcross gener- 
ations, and by study of the genetics of the natural populations them- 
selves. 



(4):26 i -174. 1963 


SYMPATRIC SPECIES 


265 



Fig. 4. Distribution of Colias hecla in North America, together with 
that of Colias meadi. The ranges of these two species do not overlap 
but are separated at their closest point by an altitudinal difference of 
4000 feet. Colias hecla is found in the mountains and plains of the 
north and Colias m.eadi only in the mountains of the south. 


266 


HOVANITZ 


/. Res. Lepid. 



Pig. 5. Distribution of Colias nastes in North America. The species 
is found only in areas north of the tree line throughout the eastern and 
central parts of its range but occurs also far to the south in the western 
region of the Rocky Mountains in the alpine tundra. 


I (4) :i6i-274, 1963 


SYMPATIUC SPECIES 


267 


It has been shown that Colias hecla and Colias nastes give all indica- 
tions of hybridizing in the wild. These indications are as extensive as 
those which are known for Colias eury theme and Colias philodice hy- 
bridizations, with the exception of a lack of the actual breeding ex- 
periments. In other words, the information is available by an analysis 
of wild populations, and character intergradation within the popula- 
tions. It has been conjectured, in other publications, that these two 
species have hybridized in a certain area, namely the Canadian eastern 
arctic, in such a way and to such an extent, that a third species, Colias 
boothi, has originated as a result of such hybridization (Hovanitz, 
1949). In the areas concerned, both parental species and the "inter- 
mediate” newly originated species all survive without the complete 
submergence of either parental species, or of the new species. The 
evidence indicates that on the fringes of the distributional area occu- 
pied by the "new species,” intermediates are present in a way suggest- 
ing introgression only, rather than the independent existence of the 
new species. This "origin of a new species by hybridization” has been 
of theoretical significance in problems of evolution for many years; 
the present example is the first clear authenticated case of this type 
of origin which can be observed in action. 

II. GEOGRAPHICAL RELATIONSHIPS BETWEEN COLIAS 
HECLA, COLIAS NASTES AND COLIAS PALAENO 

The three most "northern” Colias in the world are Colias hecla, 
Colias nastes and Colias palaeno, the most northern of these three 
being first and palaeno being least northern. Although only the first 
two appear to be involved in the natural hybridization relationship, the 
third is considered here because of its frequent sympatric geographical 
distribution. 

The world distributional ranges of these three species are shown in 
figures 1, 2, and 3. The North American ranges are shown in more 
detail in figures 4, 5 and 6. The latter three maps have been pub- 
lished previously (Hovanitz, 1951). 

Colias hecla is distributed from northern Norway and Sweden 
across the arctic shores of Asia ( fig. 1 ) , across the Bering straits, across 
Alaska and northern Canada, throughout the Canadian arctic islands 
and around all coasts of Greenland. Southern extensions of range 
occur in several places around this circumpolar range. In Scandinavia, 
the species extends part way down the Sandinavian peninsula, but it 
does not appear in more southern European locations. In Asia, besides 
the coastal arctic locations, the species exists inland in the Verkhoyansk 
area and along the Lena river valley. Farther southwards, there is much 
material indicating the possibility of its existence in almost all ranges 
of central Asia including all the area from Hindukusch, Pamir and 
Karakorum across the Himalaya to Ladakh and mountains east and 


268 


HOVANITZ 


/. Res. Lepid. 


north. The ranges extending north and eastward from the Pamir to the 
Baikal and Transbaikal completes a circle of mountains around the 
Gobi desert where some form of Colias hecla appears to live. Dots 
marking the exact locations for this area on the map were omitted 
because of the difficulty in every case to be certain of the identifications 
between the hecla group of forms and a group related to Colias meadi 
of the mountains of North America. Both of these groups appear to 
be in the same areas of central Asia, though they are separated in North 
America into contiguous regions. 

In North America, Colias hecla is found in Alaska in all areas ex- 
cept the parts southern and coastal of the main southern mountain 
masses. This includes the mountains and valleys of the interior, and 
the western and northern coastal plains. In the Rocky Mountains, the 
species is found as far south as the present location of the Alaska 
highway from Whitehorse to Ft. Nelson but there are no locations 
known south of this point in the mountains area. The species then 
extends southwards on the Rocky mountain piedmont and the valley 
of the tributaries of the Mackenzie River as far south as Nordegg (just 
west and south of Edmonton, Alberta). The range skirts the prairie 
country of central Canada near Lake Athabaska, eastward to Fort 
Churchill on the Hudson Bay. The species is found on the most north- 
ern part of the Ungava peninsula and the upper tip of Labrador, but not 
along the Labrador coast. The map (fig. 4) in addition to showing 
the distributional range of C. hecla also shows the range of C. meadi, 
the relationship of which to C. hecla is uncertain despite its close 
phenotypic appearance. 

Colias nastes has a distributional range only slightly more "southern” 
than that of Colias hecla. Actually, the ecological requirements of C. 
nastes, besides a temperature generally below 50 °F., are somewhat 
different from that for C hecla as will be seen later. Unlike C. hecla, 
Colias nastes does not exist on any part of Greenland or the most 
northern of the Canadian arctic islands. Thus, on its circumpolar 
range, there is a break of great proportions across the Atlantic which 
does not appear on the map for C. hecla. Colias nastes is found in 
Scandinava in the same general locations as Colias hecla, extending part 
way down the mountain range. From there, it extends across the 
European and the Asiatic coastal sections (presumably) across the 
Bering straits to Alaska, along the north coast of Alaska and Canada 
to Victoria Island and Baffin Island but not to the islands to the north. 
In the islands north of Europe and Asia, C, nastes is known in Novaya 
Zemlya and has been listed as having been taken in Spitzbergen. South- 
wards, the species as such is not known in Europe, unless Colias phi- 
comone of the Alps and the Pyrenees should be considered the same 
species. There seems little reason for not so considering it, in view of 
the fact that they are so similar in most respects. However, if they 
were connected in their distributional range at some earlier time, as 


(4):i6i-274. 1963 


SYMPATRIC SPECIES 


269 



Fig, 6. Distribution of the Faccmiwm-feeding Colias of North Amer- 
ica. These include Colias palaeno of the mountains and plains of the 
northwest, Colias interior of the plains in the southern part of the 
range as well as the Cascade and Appalachian Mountains, Colias pelidne 
of the eastern arctic and the northern Rocky Mountains, and Colias 
l)ehri of the alpine regions of the Sierra Nevada. 




iisiHr . j ' \ f 

' I v". iiieiii 

iii I \ ^•mn 

ww jJl ^ itaiaiai 

BEn l ^ ' i■l■l■•■t 

9MBw ■■■at fir*** aiaiaiai«n ^ 

if w k i lfc. i . f MSmF T >•«■«< >, 

fmS!tX\ \ I' i ' ME>»1^M.,^^iiaia >> 


mil I . : I ii^ fcit IVP^^k: 

a £ III J Ji/ fl L Jf laisliaK^aiaiaia 

fflwl winiiiwy*********************************?'^™ fc ■ rffii 

iiaiiaiaiaiaiaiaiaiaiaiaiaiaiaiaiaiaiaiBtttiiaiiitiaiaiaiaiaiaiaiaiaiaiaiaiaiaiaSaiaSa 

lataiaiaiaiBaaiaiaiaiaiaiaiaiBiaiaiaiiiaiaiaiaiauiBiaiaiaiaiaiaiaiaiaiaaaiaiaiBiaiafa 

laiaiaiaiaiaiaiaiBiBiaiaiaiaiaiaiaiaiaiaiaiaiaiaiBiaiaiaiaifiaiaiaiaiaiaiaiaiaiaiami 

laiBiaiaiaiBiaiaiaiaiaiaiBiaiBiaiaiaiBiiiasaiBiBiaiaiaiBiaiaiaiaiaiaiaiaiaiaiaiaiaiaie 

iami iiMi»aiJiam aiBiaiatatnatnnBiBiaiaiaiataiBinaiaiaiaiBiaMaiaaai«i«i5iSi5M«S5i5 

ia|a|ii|g||||||H|^ataiaiaiaiBiaiBiataiaiaiiiiii|I|||i|i^^aiaiaiaiaiaiaiaiaiaiStaia 

ii MWi^^^Bk amaii£a^»Biataiaiatatiri^MB^»lwikinasJllil^iiaiaiata 

ii awKy ^ t I- r^' S'* SR’IKi^Hili^EgflKiaiaii 


•2 ^ 
5j •-< 

M -C • 

U 00 

a 

^12 

<L> • 

G Of^ 
^ 3 
>» O •— » 

S5 s 

Hi 

'h3 


o 

§ 

G 

c'J 

O ^ 
.. • G 


^00 'S 

gse 

<->3 3 
^ O 
'o 

^ O 


O 

.52 N 
M.ti 

< § 

14 > 

<yi O 


-14 

’t3 
'■m' G 
G ^ 
(U 

M PQ 

(L» 


T3 ^ 

Gvo 

<SJ fNJ 


>« 


o o 
Oo 
^ON 

Sii 

(U 


5P S 

• •H 




Fig. 8. Series of Coiias, mostly identifiable as Colias boothi, showing 
by means of a graded series, nine steps in the border pattern transformation 
from the male (or hecla) pattern at the top left to the female (or nastes) 
pattern at the second to the bottom right. Grades are designated as 8 through 
4 on the left column and 3 through 0 on the right column. The specimen on 
the bottom right does not enter the series but is shown to illustrate a 
weak pattern development on a grade 0 male. All specimens shown are 
males and all are from Coppermine, Northwest Territories, Canada, July 12- 
19, 1947, W. Hovanitz coll, except: (1) 2rd from top on right column: 
Repulse Bay, Northwest Territories, Canada. July 21, 1950. P. F. Brugge- 
mann. (2) 5th from top (last one) in right column. Repulse Bay, July 15, 
1950. P. F. Bruggemann. 






272 


HOVANITZ 


/. Res. Lepid. 


during the Pleistocene ice age, many thousands of years of isolation 
since that time has brought differences between them and earned a 
species distinction. Southward in central Asia in the ring of mountains 
surrounding the Gobi desert, including the Himalayas there is a string 
of races all of which appear to be specifically related to C. nastes; the 
main form here is known as C. cocandica. The map for C. nastes 
therefore has quite a similarity to the map of C. hecla. It can be seen 
however that in the area of Asia west of Sakhalien Island, C. nastes 
occurs considerably south of the area of C. hecla. 

In North America, Colias nastes occurs also farther south than C. 
hecla. The species extends southwards in the Rocky mountains from 
the mountains of Alaska and the Yukon Territory to the borders of 
the United States in British Columbia and Alberta. In Alberta, C. 
nastes occurs at an elevation of 7500 feet and up while C. hecla is 
found at about 3-4000 feet in the piedmont. In these areas, C. nastes 
is flying in late July and August while C. hecla at the lower elevations 
is flying in May. C. nastes does not extend into the lowland valley areas 
of the Mackenzie tributaries as does C. hecla but instead is found only 
to the north of the tree line from the mouth of the Mackenzie to the 
area of Ft. Churchill on the Hudson Bay. It is found on the Belcher 
islands in Hudson Bay. It is found in the northern part of the Ungava 
Peninsula and then southeastwards along the Labrador coast half way 
down. The fost northerly known locality is on the northern coast of 
Baffin island. These locations may be observed in relation to one 
another on the map (fig. 5). 

Though Colias palaeno is not involved in the specific study here 
being analyzed, it was thought that it might be so involved and therefore, 
its geographical distribution is here given together with the two pre- 
ceding species. This species has a generally more southern distribu- 
tional range than either Colias hecla or Colias nastes as can be seen by 
study of the world map (fig. 3). The species is found in the Alps of 
Europe, in a number of locations in the territory intervening between 
the Alps and the Scandinavian mountains, and northwards and east- 
wards. It does not exist far south of Lake Baikal in central Asia but 
does extend southwards into Northern Korea, on the island of Hondo 
in Japan, Sakhalien Island, and Kamtchatka Peninsula. The species is 
not known on the islands north of Europe and Asia. In North Amer- 
ica, the species is known in Alaska south of the Brooks range and north 
of the main southern ranges which skirt the coast. It reaches the coast 
of Northern America only at the mouth of the Mackenzie river but 
extends southwards along the route of the Alaska highway to a point 
just north of Ft. St. John and thence eastwards to the Riding Mountains 
of Manitoba and northwards to the tip of the Ungava Peninsula. Its 
range is limited southwards and eastwards by contact with Colias pelidne 
(also called interior). For details on this contact zone, the map of 
North America is more precise (fig. 6). 


1963 


SYMPATRIC SPECIES 


273 


IIL CHARACTERISTIC DIFFERENCES BETWEEN COLIAS 
HECLA, COLIAS NASTES AND COLIAS PALAENO 

Colias hecla may be distinguished readily from Colias nastes by 
the fact that it is always orange in wing color (fig. 7), while Colias 
nastes is pale yellow. Colias palaeno is more generally a brighter lemon 
yellow (fig. 7). 

A second character that separates hecla from nastes but not from 
palaeno is a male-female dimorphism of the pattern. The male pattern 
is a single solid band of black on the outer edges of the fore and hind 
wings. This appears on the wings of the males of both hecla and 
palaeno (fig. 7). The female pattern on the other hand dif feres from 
this in a way that might be described as a series of dots in the border 
band, or in what might be the band when it is all present. This is 
typically shown for the females of hecla and nastes on figure 7. The 
band of the female of Colias palaeno is so reduced that the dots show 
only slightly in that species. The male of Colias nastes differs from 
any other North American species of Colias in that it also shows a 
series of dots in the border band, much like the female. In some 
cases, it is very difficult to distinguish the males from the females by 
general appearances. 

Other differences in habit, larval food plant and ecological prefer- 
ences will be discussed later in this series. 

Intergradation between Colias nastes and Colias hecla as would be 
caused by hybridization, the crossing of Fi to obtain the F 2 segregation 
and backcrossing should give a series of intermediate products with 
all intergradations from the one parental type to the other. In addi- 
tion there should be produced, if the genetic segregation truly assorts 
itself with slight effects of genetic linkage, into truly divergent types 
such as a fully orange male with the female border band, a type found 
nowhere under natural conditions in the ordinary range of the species. 
Another type of extreme divergence which would be found only under 
such conditions of gene assortment would be a yellow form with the 
typically male border pattern. All these types have been found in 
areas of presumed hybridization of these two species, Colias hecla and 
Colias nastes. A series of variations ranging from the male border band 
to the female border band in a series of nine steps (grades) is shown 
in figure 8. A similar graded series of nine steps for the orange pigment 
on the upper side of the wings is shown in figure 9. These series are 
to be used to illustrate the analysis of various populations of Colias 
from diverse regions in the arctic. 


274 


HOVANITZ 


/. Res. Lepid. 


LITERATURE CITED (I, II, III) 

HOVANITZ, WILLIAM. 1943. Hybridization and seasonal segregation in 
two races of a butterfly occurring together in two localities. Biol. Bull. 
85: 44-51. 

— — — ■ 1944a. The ecological significance of the color phases of Colias 

chrysotheme in North America. Ecology 25: 45-60. 

— — 1944b. Genetic data on the two races of Colias chrysotheme in North 

America and on a white form occurring in each. Genetics 29: 1-30. 

— — -- 1948. Ecological segregation of interfertile species of Colias. Ecology 
29: 461-469. 

1949. Increased variability in populations following natural hybridiz- 
ation. In ’'Genetics, Paleontology and Evolution,” Princeton University 
Press, pp. 339-355. 

— - — - 1951. The biology of Colias butterflies. I. The distribution of the 
North American species. Wasmann J. Biol. 8: 49.75. 

1956. Hybridization and species blending in the butterfly genus 

Colias. Proc. XIV Int. Congr. Zool. Copenhagen, 1953, p. 140. 

SETTE, O. E. 1962. Variation in the silvering of Argynnis {Speyeria) callippe 
in the interior m*ountain area of southcentral California. /. Res. Lepid. 
1 ( 1 ): 3 - 20 . 


(to be continued) 


Journal of Research on the Lepidoptera 1 (4) :275-279, 1963 

1140 W. Orange Grove Ave., Arcadia, California, U.S,A. 

© Copyright 196 } 


A METHOD FOR BREEDING PIERIS NAPI 
AND PIERIS BRYONIAE 

BJORN PETERSEN 

Zoological Institute, University of Lund, Lund, Sweden 

Since the year 1942 the author has bred Pieris napi L. and 
Pieris hryoniae Ochs, over a number of years. This breeding was carried 
out mainly in order to obtain information on the genetical, ecological, 
and taxonomical relationships between the two forms (cf. Petersen 
1963 , in press, and literature cited there). This paper will deal with 
the breeding methods used and some results obtained in this connection. 

MATING 

For copulation males and females were put together in cylindrical 
cages made of cotton net with a bottom of treetex. The cages were 
2-4 dm high and had a diameter of 2-3 dm. In such a cage 2-4 females 
and a slightly larger number of males were placed. The cages were 
hung up in a window facing towards the south. Quite soon it was 
obvious that mating propensity was higher for napi in sunshine and 
for hryoniae in cloudy weather ( cf . table 1 ) . Mating propensity means 
in this case the frequency of maxing of one sex compared with the 
same sex of the other species. The difference is statistically significant 
(X^=l6.37 with Yates’ correction; P<0.001). Muller and Kautz 
(1938 p. 15) state that sunshine is essential for the copulation of 
hryoniae. In nature hryoniae, like napi, copulates in sunshine. The 
experiments indicate only different ecological amplitudes for the copula- 
tion of the two species. 

Matings under laboratory conditions were fairly easily obtained, not 
only by the author in Sweden, but also by Bowden and Easton in Eng- 
land (Bowden 1953, 1956, 1957, Bowden and Easton 1955). Ento- 
mologists in Central Europe, on the other hand, have had a number of 
difficulties in obtaining such matings. Some few experiments on hybrid- 
ization have been reported by Fischer (1924, 1925) and Kautz (Muller 
and Kautz 1938 p. 159-161). The lack of success in many of the 
Central European experiments may be due to too high temperature 
during sunshine in Central Europe. The temperature in the windows 
used by the author, when the experiments were made in April and 
May, was usually about 24°-27°C. In the sunshine of Central European 
laboratories the temperature probably is higher, especially as the ex- 
periments usually are carried out during the summer months. 


275 


276 


PETERSEN 


/. Res. Lepid. 


TABLE 1, Number of matings (f) of animals tested on 
mating propensity (m) under different conditions. Sim- 
plified from Petersen and Tenow, 1954, p, 182, 



sunshine 

cloudy 

weather 


f 

m 

f 

m 

napi 

31 

0,84 

1 

0,14 

bryoniae 

6 

0,16 

6 

0,86 

napi 

22 

0.76 

2 

0,29 

bryoniae 

7 

0,24 

5 

0,71 


TABLE 2, Survivals from growing P* bryoniae from 
Abisko, northern Sweden, 


temperature 

number of 

eggs 

larvae 

pupae 

40 

10 

0 

0 

80 

10 

3 

0 

13° 

10 

9 

6 

16° 

10 

10 

9 

20° 

20 

20 

11 

28° 

10 

7 

0 

0 

CM 

00 

22 

14 

0 

0 

=i- 

CO 

5 

0 

0 


(4)s275-.i79. 1963 


BREEDING PIERIS NAPI 


277 


EGG-LAYING 

After copulation, the female was transferred into a cage with some 
flowers to feed on and leaves of a species of the family Cruciferae for 
egg-laying. P. bryoniae in nature lays the eggs almost exclusively on a 
single plant species, Biscutella laevigata. In one cage experiment a 
female preferred some other plants to Biscutella, about 80 eggs being 
laid on them and none on Biscutella ( Petersen l.c. ) . Thus, every plant 
appropriate for the egg-laying of napi seems to be appropriate for 
bryoniae. However, most plants of families other than Cruciferae as 
well as some few species of this family are not eaten at all by the larvae. 


TABLE 3. Results from breeding napi - bryoniae from four 
localities in Sweden, 1. = number of newly hatched larvae, 
p, s number of pupae. 


temp. 

Scania 

Uppsala 

Murjek 

Abisko 

1, 

P* 

% 

1, 

P* 

% 

1, 

P« 

% 

1, 

P* 

% 

11®-12° 

30 

11 

37 




10 

2 

20 

15 

0 

0 

15®-16° 

22 

7 

32 

25 

19 

76 




11 

1 

9 

16^-17° 

85 

46 

54 

5 

5 

100 

4 

3 

75 

24 

12 

50 

190 - 20 ° 

153 

119 

78 

45 

26 

58 

17 

8 

47 

33 

19 

58 

220-23° 

43 

29 

67 




5 

2 

40 

36 

11 

31 

250-26° 

92 

47 

51 

5 

4 

80 








STERILITY 

In many broods of the species as well as of their hybrids, the eggs 
do not develop at all or only partly. Sterility appears with almost the 
same frequency in both types of crosses, 6 broods of 30 in the pure 
species, 6 of 28 in the hybrid crosses. Sterility is slightly more frequent 
among Bowden’s Fi hybrids and backcrosses, but the difference is not 
significant. The F 2 and F 3 crosses, on the other hand, have a significantly 
higher frequency of sterility, when compared with the Fi crosses and 
back-crosses of Bowden (x^ = 5.94 with Yates’ correction, 0.02 >P 
> 0 . 01 ). 


278 


PETERSEN 


/. Res. Lcpid. 


TEMPERATURE 

The influence of temperature on the breeding results was investi- 
gated in two series of experiments. In the first series, material from 
Abisko in northern Sweden was used. P. bryoniae flies here from 400 m 
up to 800 m in the subalpine and the lower parts of the alpine regions. 
The breeding results are shown in table 2. The optimal constant tem- 
perature for eggs and larvae was found to lie between 13° and 20° C. 
As the hibernating pupae require a rather low temperature the experi- 
ments were stopped at pupation. 

In the second series, material from four Swedish localities was used 
( cf. table 3 ) . The Abisko material developed fairly successfully from 
newly hatched larvae to pupation at 16° -23° C. The material from 
Abisko was perhaps a little weaker than that in the experiment already 
described, as the eggs had to be sent by mail. The material from south- 
ern Sweden (Scania) developed at temperatures varying from 11° to 
26° C. The adaptation to different climates is obviously not strong 
enough to show up in experiments on such a small scale, especially near 
the upper and lower limits of temperature. 

DISEASES 

A successful breeding is dependent, not only on appropriate food 
and temperature, but also on the health of the animals. Virus diseases 
often cause the blackening and death of the larvae. The cultures started 
first usually develop well, only some of the latest-developing larvae 
being affected. In later cultures the disease spreads like a pest, killing 
the larvae in great numbers. The results of table 3 may therefore better 
be expressed as different resistance to the disease at different tempera- 
tures, or perhaps as varying virulence of the virus, than as a direct effect 
of the temperature. 

Most years, the larvae were kept in glass jars on cut leaves. As the 
disease seemed to spread more rapidly on wet leaves, it was decided 
to start breeding on living plants growing in flower pots. The results 
were promising in that the frequency of virus disease became very low. 
The fecundity of the animals, on the other hand, was not very high 
during this last year of more extensive breeding. It has been stated 
by several breeders of Pieris napi that it is impossible to breed many 
generations without crossing with fresh material taken in nature. The 
animals used belonged to the fourth generation in captivity and were 
unfortunately the only ones available at the time. 

The best way of breeding Pieris napi has still to be investigated. ' 
Breeding at 20° C on a living plant of the family Cruciferae, which 
is easy to keep in flower pots, will probably give better results than 
any so far known. Biscutella laevigata, which grows as a weed in the 
gardens of Italy, will perhaps be the best food-plant for P. bryoniae. 


( 4 ): 275 - 279 . 19^3 


BREEDING PIERIS NAPI 


279 


LITERATURE CITED 

BOWDEN, S. R. 1953. Timing of Imaginal Development in Male and 
Female Hybrid Pieridae. (Lep.) Entomologist, 86, No. 11: 255-264. 

1956. Hybrids within the European Pieris napi L. Species 

Group. (Lep., Pieridae). Proc. South London Ent. Nat. Hist. Soc., 
1954-55: 135-159. 

— , 1957. Diapause in Female Hybrids: Pieris napi adalwinda 

and Related Subspecies (Lep.) Entomologist, 90, No. 1133: 247-254, 

273-282. 

BOWDEN, S. R. and N. T. EASTON. 1955. Diapause and Death: Further 
Observations on Imaginal Development in Pieris Hybrids (Lep.) Ento- 
mologist, 88, No. 1108: 174-178, 204-210. 

FISCHER, E. 1924. Ober die Zweibriitigkeit der P. bryoniae O. Mitt. 
Munchen. Ent. Ges., 14:8. 

— . 1925. Neue Zuchtergebnisse bei Pieriden. Mitt. Schweiz. 

Ent. Ges., 13- 

MULLER, L. and H. KAUTZ. 1938. Pieris bryoniae O. and Pieris napi 
L. Wien. 1- 189. 

PETERSEN, B. 1963. Breakdown of differentiation between Pieris napi L. 

and Pieris bryoniae Ochs, and its causes. Zool. bidr. Uppsala, 35: in press. 
PETERSEN, B. and O. TENOW. 1954. Studien am Rapsweissling und 
Bergweissling (Pieris napi und Pieris bryoniae). Isolation und Paarungs- 
biologie. Zool. bidr. Uppsala, 30: 169-198. 



Journal of Research on the Lepidoptera 


1 ( 4 ) : 281 - 300 , 1963 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 


AN ANALYSIS OF THE NORTH AMERICAN SPECIES 
OF THE GENUS CALLOPHRYS 

J. W. TILDEN 

San Jose State College, Sort Jose, California 


INTRODUCTION 

EXAMINATION OF CERTAIN SPECIMENS of Callophrys from mon- 
tane habitats in California and Oregon indicate that these specimens 
belong to a cluster distinct from others so far described, and not fitt- 
ing well with any known description of North American Callophrys. 
The purpose of this paper is to discuss certain characteristics of North 
American Callophrys, and to provide a name for the previously un- 
recognized series of populations. This name is needed for two forth- 
coming faunistic papers. 

TAXONOMIC HISTORY OF CALLOPHRYS 

Linnaeus (1758) described P (apilio) P {lebeji) rubi as follows: 
‘Rubi. 154. P. P. alls dentato-subcaudatis; supra fuscis, subtus viri- 
dibus.” 

Billberg (1820) proposes the genus Callophrys with three groups, 
based on three tails, two tails and one tail. In the one-tailed group only 
rubi is listed. 

Scudder (1875), selected rubi as the type of Callophrys as follows: 
'T820, Billb., Enum. Ins. 80: Vulcanus, rubi, and a Ms. species. Rubi 
may be taken as the type.” 

Until recently Callophrys has been limited to those few hairstreaks 
that matched rubi in every respect. The palaearctic names, besides rubi 
itself, include borealis KruL, polaris Moschl., fervida Stgr., sibirica 
Ruhl and suaveola Stgr., omitting abberations that have been named. 
All of these have been considered both as full species and as subspecies 
or variants of rubi. Of these only rubi, as type of genus, will be men- 
tioned further, "f 

Ziegler (I960), combined the groups placed under Mitoura Scud., 
Incisalia Scud., Sandia Clench & Ehrlich, and Callophrys Billb., under 
the oldest name, Callophrys, as subgenera. This move had been pre- 
sented in abbreviated form previously by Ehrlich and Clench (I960) 
in the paper describing Sandia mcfarlandi. Clench, in Ehrlich and 
Ehrlich (1961) used a similar arrangement but several genera are 


281 


282 


TILDEN 


/. Res. Lepid. 


proposed for groups of hairstreaks outside the scope of this paper, 
which is concerned only with the members of the presently considered 
subgenus Callophrys as limited by Scudder’s designation of rubi as the 
type. 

THE AMERICAN SPECIES OF THE SUBGENUS CALLOPHRYS 

All species of the subgenus Callophrys resemble one another very 
closely, and can be separated with any degree of assurance only by 
considerable effort. There always remain specimens the position of 
which is a matter of opinion in the light of present knowledge. There 
are two revisions of the group. Barnes and McDunnough (1923), 
grouped the species in a manner superior to previous treatments and 
described three subspecies. Clench (1944) revised the genus (as it 
was then considered) proposing one subspecies. 

At present at least ten names are involved in the North American 
Callophrys (s. str.), and these have been regarded at one time or 
another as representing six distinct species. Characters that have been 
used to distinguish the various species have been color markings and 
wing shape. The male terminalia are exceedingly similar in all the 
named entities. This is not to say that differences may not exist, but 
so far they are not apparent. The present author has also dissected the 
female genitalia of the North American species and the European 
rubi. These studies have not resulted in any conclusions in time for 
the present paper. 

An attempt has been made to examine all parts of the external 
surface of the body. Small differences may be detected in the palpi, 
antennae, facial hairs or tuft, and in wing shape, stigma and color 
markings. The results of these comparisons are given in Table 1. 

It seems pertinent to include here a discussion of the problem of 
the identity of dumetorum Bdv. Boisduval ( 1852) writes (free tran- 
slation from the original French) that "this Theda resembles our 
rubi in every respect and is most likely only a local variety of that 
species.” Close examination of rubi in comparison to specimens usually 
considered to represent dumetorum Bdv. shows a number of minor 
differences, but without doubt the two are closely related. Certain 
problems arise in deciding what would properly be considered as 
dumetorum Bdv. Clench (1955, op. cit, p. 220) gives a description 
of what he at that time regarded as dumetorum. The description 
suggests that he is describing the insect for which a new name will be 
proposed in this paper. 

The Boisduval description could fit any of several named segre- 
gates. The figure by Oberthur (1913) shows a gray insect with over- 
scaling of a cast of green not usual in the subgenus, and with a rather 
more complete macular band than one expects in insects that pass 
for dumetorum in collections. W. D. Field, of the Smithsonian In- 


(4):28i-}00, 196J 


NORTH AMERICAN SPECIES 


283 


sitution, has kindly given me a detailed comparison of the insect 
figured by Oberthur, and which is usually regarded as the type of 
dumetomm. Whether or not this specimen is actually the one on 
which Boisduval based his description is not certain; this does not 
seem to be stated explicitly. Field- informs me that Oberthur’s figure 
is a faithful representation of the supposed type specimen. This speci- 
men is a gray female with the fringes white or pale-tipped with a con- 
spicuous macular band and no antennae. 

This female specimen, by virtue of being illustrated by Oberthur, 
may need to be regarded as a lectotype, the illustration being one 
method of designation, as was suggested by Field (in litt), A possi- 
bility exists that it may indeed be the specimen described by Boisduval, 
but this point can not be settled in light of present knowledge. 

Neither Boisduval’s description nor Oberthur’s illustration seem 
to fit exactly any presently known population of Callophrys. Either 
dumetomm was described from unusual specimen, or from some 
since unlocated population, or (most likely) neither the illustration 
nor the description are completely recognizable in relation to some 
well-known population. Of the specimens compared with the type, 
there seem to be fewer discrepancies with the lowland population 
most commonly considered to be dumetomm, except that the type is 
a gray female while the females of the lowland population tend to 
be brown or fulvous. However, gray females are not entirely un- 
known. The macular band is more than usually well-developed in 
the "type,” but the band is variable in all species, and specimens 
from the foothills of the Sierra Nevada are known with complete 
macular band. The choice here made is to continue to use the name 
dumetomm for the cismontane lowland population of California, while 
retaining the option of changing this opinion should future findings 
make such a change desirable. 

The exact locality where Lorquin collected his original specimen, 
subsequently named dumetomm by Boisduval, is partially in doubt, 
as is frequently the case with the Lorquin-Boisduval specimens. 
Species described in 1852 were from Lorquin’s earlier collecting. In 
some of the descriptions there are phrases such as "Mountains of the 
Juba,” suggesting that the specimens were taken near the mining 
operations along the Yuba River. A reading of Lorquin’s adventures 
in California brings the same conclusion; that he was in the Sierran 
mining region prior to 1852. This is presumptive evidence, but 
certainly better than no inference at all. Specimens of Callophrys from 
the Sierra Nevada foothill localities relate to the populations from 
coastal and southern California rather than to those of the high Sierra 
Nevada. Here again the evidence favors use of the name dumetomm 
for the low elevation insects. 

Should incontrovertible evidence subsequently be found, that this 


284 


TILDEN 


/. Res. Lepid. 


Character 

dumetorum auct. 

apama 

viridis 

wing fringes 

fuscous basally; 

pale tipped; not 
clear white 

dark fuscous basally; 
pale tipped, dark 
scales mixed. 

fuscous; the tips 
mixed pale and 
fuscous scales 

fore wing apex 

obtuse-angled 

obtuse-angled 

obtuse-angled 

forewing, outer 
margin 

oval in female; 
straighter in male 
and indented at Cu 2 

oval in female; 
male slightly pro- 
duced between Cu]^ 
and 

curved to or M ; 

then nearly straight; 
tornus slightly 
incurved 

hindwing, outer 
margin 

crenate in spaces 
Cuj^, Cu 2 and 2nd A. 
(indentations shallow) 

small crenation in 
space Cu 2 ; a deeper 
one in space 2nd A 

hind wing appearing 
more than usually 
quadrate; crenations 
very slight 

color of fore- 
wing costa 
below 

fulvous, more marked 
in female 

pale brown to dull 
fulvous; concolorous 
with other brown 
areas of wing 

rich fulvous in both 

sexes 

forewing below 

invaded by gray or 
fulvous from anal 
margin to at least 
vein M]^ 

anal margin gray; 
disc invaded by rich 
fulvous at least to 
vein M]^ 

gray confined to anal 
cells only 

vein-tips of 
hind wing below 

vein tips and marginal 
line usually rusty- 
brown scaled 

narrow but complete 
marginal line, out- 
brown, mesially 
black, inwardly 
white; vein tips dark 

brown scales at vein- 
tips absent or nearly 

so 

Macular band 
(lower surfaces 
of wings) 

usually reduced to 

3 spots; seldom com- 
plete; macules in- 
wardly brown, then 
white 

hindwing: complete, 
spots in Cuj^ 6i Cu 2 
displaced out. Tri- 
colored, 4-5 spots 
forewing 

complete on both wings 
(though narrow) to re- 
duced; mesial brown 
scaling reduced 

shade of green 
below 

grass green to 
golden green 

rather dull green, 
brightened by ad- 
mixture of fulvous 
scales 

deep, frequently 
bluish, green. 

scaling of 
labial palpi 

black above, mixed 
black 6c white below; 
may have few green 
scales near base 

hairs and scaling 
smooth; black above 
and below; white 
scales laterally 

grizzled black and' 

white throughout; 
third segment very 
pointed; scaling 
sparse, green scales 
basally 

facial hairs 
(facial tuft) 

erect or slightly 
proclinate; thicker 
laterally; black with 
green scales at bases 

hairs markedly pro- 
clinate, thicker 
laterally; basal 
green scaling 

erect or slightly 
proclinate; thicker 
laterally; black or 
gray; green scales 
at base 

forewing 
stigma of male 

small, ovate, usually 
gray and lighter than 
ground color 

ovate to oval; 
gray, slightly 
lighter than ground 

small, ovate, dis- 
tinctly paler than 
ground color, or 
concolorous 

antennal annuli 
(white rings) 

usually 15 (seldom 

16) 

usually 15 (seldom 

16) 

14-17 (av. 16 in 34 
specimens) ; antennae 
pale above (unique) 

general facies as 
seen from above 

male uniform fuscous 
with pale- tipped 
fringes; female brown 
with fulvous discs 
and pale- tipped fringes 

male uniform dark 
fuscous; female 
same with very 
large fulvous discal 
areas 

gray 

insect with nearly 
concolorous fringes; 
females usually gray 
(very seldom partly 
brown on disc) 


TABLE 1 


CHARACTER COMPARISON OF NORTH AMERICAN CALLOPHRYS (S.STR.) 


(4);28 i -300, 1963 


NORTH AMERICAN SPECIES 


285 


affinis 

sheridani 

corns tockl 

lemberti n. sp. 

basally fuscous; 

pale- tipped to 
white tipped 

bases darker than 
wing; tips mixed 
white & fuscous 

bases darker than 
wing; tips white , 
contrasting 

basal scales mixed ) 

brown and gray; tips 
snow white (usually); 

approx, right- 
angled, tip rounded 

acute, the forewing 
trlgonate 

rectangular; effect of 
short sharp tip 

pointed, nearly as in 
sheridani! 

curved to , 

thence nearly 
straight 'to tornus 

alight curve to 

or M 2 , then straight 
to tornus or 
slightly concave 

quite evenly curved, 

the curvature 
slight 

curved to M 3 ^ 

slightly indented 
between Cuj^ & Cu 2 

quadrate, nearly 
rounded; crenations 

slight or obsolete 

rounded; crenations 
scarcely visible 

longest between M 3 
& Cu^; crenations 
evident but minute 

rounded, crenations 
slight, between CU 2 
& 2nd A one is 
evident 

gray to clay- 
colored, not 
contrasting 

blackish; darker 
than rest of wing 

dark in most males; 
concolorous to pale 
fulvous in females 

narrowly brown or 
fulvous; contrasting 

gray from anal 
margin to Cu ; 
general wing 
surface green 

dark gray to 
vein Cu 2 

gray area extensive, 
to M 2 (one female) on 
to across wing to 
costa (one male) 

gray confined to anal 
cells; even here some 
green overscaling 

slight brown at 
each vein- tip; 
terminal line 
not differentiated 

spots at vein tips 
not evident; term- 
inal line black 

dark vein tips not 
evident; terminal 
line black, narrowly 
white Inwardly 

usually slight dark 
points at vein tips 
on both pairs of 
wings; approaches 
checkering 

obsolete, or one 
spot in cell Cu 2 

complete; straight; 
white; edged within 
and without by black 

band complete but of 
separate spots; bowed 
out at cell Cu 2 ;mac- 
ules white, black 
inwardly . 

^ complete band to 
obsolete of discrete 
white spots black 
inwardly, narrow; 
obscure on forewing 

uniform pale 
yellow- green 

deep dark green scales 
mixed with dark gray 
scales about equally 

deep dark green, the 
veins slightly con- 
trasting dark 

green scaling thin, 
luniform, bright pale 
‘green, the undercolor 
showing through 

palpi slender 
and thin scaled; 
mixed black and 
gray all over 

dark above; scaling 
below dense; mixed 
black and light 
gray (effect dark) 

palpi slender, 
pointed; scaling 
sparse; dark above, 
mixed black & white 
below 

palpi slightly darker 
above but mixed white 
and black scales 
throughout; third 
joint darker 

facial hairs 
light gray, 
sparse, procumbent 

dark, dense, slight- 
ly proclinate; 
underscaling pris- 
matic, not green 

tuft dense, coarse, 
b^ack, proclinate; 
subscaling covered 
or obscure 

hairs sparse, fine 
dark, sub-erect; 
green sub-scaling 
prominent 

dark to black, 
sharply con- 
trasting 

stigma usually 
nearly concolorous 
with wing 

small, slightly pale 
to concolorous and 
scarcely discernible 

stigma small, sub- 
triangular, con- 
colorous to pale; 
seldom dark 

17 

av. 17 (16-18) 

15-16 

16 (17 on one 
specimen) 

fulvous to bright 
rufous with dark ter- 
minal line & pale 
fringes 

dark gray with black 
terminal line and 
white fringes; sexes 
alike in color 

gray insect with 
dark terminal line 
and white fringes; 
sexes concolorous 

mouse gray smooth- 
scaled insect with 
white fringes; females 
concolorous or dull 
brown (fuscous) 


TABLE 1 (continued) 


286 


TILDEN 


/. Res, Lepid. 


rubi 

Incisalia augustinus 

Mitoura siva 

Sandia mcfarlandi 





basally dark; tips 
with a few pale 
hairs 

mixed brown & gray 
basally; tips sordid 
gray; dark at vein- 
tips (checkered! 

forewings fuscous; 
hindwings pale- tipped 

basally brown and 
fuscous mixed; tips 
white 

tip quadratCji bluntly 

pointed 

tip rectangular or 

nearly acute 

rather acute 

obtuse 

rounded to thence 

nearly straight to 
tornus male indented 
at Cu 2 

rather evenly curved 

but slightly flatter 
before tornus 

quite evenly rounded 

slightly flattened 
before tornus 

rounded 

two well-marked 
crenations at cells 
Cu 2 and 2nd A. 

at least a suggestion 
of a crenation between 
each pair of veins 

tailed at Cuj^ & Cu 2 ; 
not crenate 

tornal crenation 
evident, slight 

broadly fulvous 
in female; narrowly 
so or gray in male 

concolorous with wing 

slightly fulvous, not 
greatly contrasting 

lemon yellow to 
apricot 

gray confined to anal 
cells 

no contrast 

fulvous invasion from 
anal margin almost to 
costa 

yellow to apricot 
shade across entire 
wing 

vein- tips brown; 
terminal line warmi 
brown in female, 
less so in male 

terminal line dark 
brown to black; veins 
not contrasting 

complex pattern of 
black, white and 
rufous overscaling 

terminal line black, 
invading vein tips 

of small separated 
narrow white spots; 
nearly complete to 
obsolete 

complete band of 
small round dark 
spots; dark basal 
shade 

complete, irregular 
from inward out of 
bands of brown, 
black & white 

narrow, complete, 
white flanked each 
side with black 

grass green- 
suggests dumetorum 

ground color 

pale vinaceous brown 

smooth pale green 
scaling where other 
patterns are not 
evident 

basically a luminous 
yellow- green 

mostly dark with 
white scales on 
sides; green scales 
usually evident 

mixed black & white 
scales; darker above 

black above and at 
tips; basally, white 
overscaling 

dark above and at 
tips; white scaling 
below and laterally; 
rather long 

sparse, dark, mostly 
lateral; median 
green sub-scaling 
prominent 

dense dark rich 
brown, procumbent; 
sub-scaling covered 

very sparse, pro- 
cumbent; iridescent 
underscaling very 
visible 

hairs fine, short 
& erect; mixed 
black & white; 
subscaling dark 

elongate-oval ; 
androconia rather 
rough 

very long oval, 
androconia small; 
black to pale 

elongate-oval ; 
dark to pale 

elongate, about 

3 times as long 
as wide 

16 

18 

1 7 , narrow and 
clear-cut 

15-16; antennae very 
short, each segment 

gray with nearly 
concolorous fringes 
(male) ; dark brown 
with concolorous 
fringes (female) 

dark brown with 
checkered fringes 
(male) ; female 
lighter richer brown 

dark brown Insect 
with fulvous on discs 
and two dark dots 
near hind wing tornus; 
female more fulvous 

brown insect with 
fulvous discs & 
white fringes 


TABLE 1 (continued) 


I (4) :28i.300, 196} 


NORTH AMERICAN SPECIES 


287 


position is untenable and that the population herinafter described as 
unrecognized is actually true dumetorum, the populations now con- 
considered as dumetorum and its variants must then become known 
as perplexa B. & McD., as the oldest available name definitely assigned 
to this coastal insect. 

The notations in Table 1 are of necessity short. In some cases 
amplification is given in the species discussions. The characters used 
in Table 1 vary in the samples from the different populations, but 
the degrees of quality given are averages. In all cases the samples were 
adequate (twenty-five or more specimens) to large (fifty to one 
hundred specimens) except for af finis and comstocki. Of the latter, 
only four specimens were available for close study, and some fifteen 
specimens were examined in all. 

For comparison, one species each of the subgenera Incisalia, Mitoura 
and Sandia (which contains a single species) are included and com- 
parison will show that most of the characters used are specific 
rather than generic or subgeneric. The subgenus Narnia was excluded 
from lack of material, and the subgenus Cyanophrys because it enters 
our fauna only along the extreme southern border. 

Clench (in Ehrlich & Ehrlich) defines Callophrys (s. lat.) and 
its subgenera primarily on genitalic characters. The subgenus Callophrys 
includes species with valvae not capped; the cornuti of the aedeagus 
slender, not spatulate; the scent-pad (stigma) well-developed, and 
the labial palpi about IV 2 times as long as the vertical eye diameter. 
(This last character is shared with the other subgenera except for 
sandia, the palpi of which are about twice as long as the vertical 
diameter of the eye). The scales of the stigma are entire and with 
rounded ends and the hind wing is not tailed. 

In addition may be mentioned the tornal "tab” of the hind wings, 
shared with Incisalia; the reduction of the hind wing marginal crenula- 
tions, restricted usually to the last three cells (much more extensive 
in Incisalia); the usually even green overscaling of the inferior sur- 
faces, typical of the subgenus, and the complete lack of the thecla 
spot. The markings below are restricted to the submarginal band 
(macular band as frequently stated), which is often reduced or even 
absent, even in a single species. There is no other evident ornamenta- 
tion. 

In the following analyses of the species, only the original citation 
is given. 


Callophrys rubi ( L. ) 

Syst. Nat. 10th Ed. 1:483, No. 154:1758 
Palaearctic. Range Europe and Asia. By some considered the only 
valid palaearctic species. Closely resembles dumetorum as considered 
here, but the green darker, the female less broadly fulvous, the wing 


288 


TILDHN 


/. Res. Lepid. 


fringes less contrasting, the stigma more elongate, and the under sur- 
face of the forewing not greatly invaded by gray or brown. In all 
specimens examined, the terminal line of the wings is much more 
evident. 


Gallop hrys dumetomm (Bdv.) 

Ann. Ent. Soc. France (2) 10:291:1852 

A fuscous (male) or usually broadly fulvous (female) insect with 
pale-tipped but not white fringes; hind wing with three crenations, 
terminal line not strongly contrasting. Green below warm grading to 
yellowish green in southern specimens, nearly distinctive. Forewing 
deeply invaded by gray (male) or tan (some females) at least to vein 
Ml and frequently (subspecies perplexa) to forewing costa. Veins 
of secondaries below dark tipped, the border usually clouded with 
brown scales. Whether or not this species is correctly identified as 
dumetomm, it is one of the distinctive entities of the subgenus. The 
invasion of the forewing by gray or brown seems diagnostic in its 
range. The subspecies perplexa B. & McD. (1923) is more yellowish- 
green below, the band reduced or obsolete (usually) and with the fore- 
wing invasion frequently extending to the costa. However, reduction 
of the macular band occurs in all populations of dumetomm (as well 
as in nearly all other populations of Callophrys) and this character 
must be used with caution. The macules of the band are bicolored, 
brown inwardly, outwardly white. 

Callophrys apama ( Edw. ) 

Papilio 2:137:1882 

A very dark fuscous (male) or fuscous with broad fulvous discs 
(female) insect with fringes slightly pale-tipped, not notably contrast- 
ing. Green below rather dull but mixed with fulvous scales, giving 
a superficial appearance of being lighter than is really the case. Macular 
band (nominate apama) complete, the macules in spaces Cui and 
Cug displaced outwardly, the macules of the band tricolored, inwardly 
brown, mesially black and distally white. Facial tuft peculiar, sparse, 
rather light in color and the hairs markedly proclinate in all of the 
specimens examined. The name homoperplexa B. & McD. (op. cit., p. 
68) was given to specimens such as those from Colorado, in which the 
macular band tends to become obsolete. The shade of green and the 
other characters remain similar. 

Callophrys viridis ( Edw. ) 

Though usually considered a synonym of dumetorum, viridis appears 
to be as distinct as most of the species, and was ressurected from 
synonomy by Clench (1944). It is a dark gray insect in both sexes 
when fresh (old specimens fade to a lighter gray), the fringes pale- 


(4):28 i -500, 1963 


NORTH AMERICAN SPECIES 


289 



Fig. 1. Callophrys. Upper side to left; lower side to right. Callophrys 
mbi L. Near Hannover, Germany, 12:IV.52. G. Hesselbarth. 

Callophrys dumetorum Bdv. El Portal, Mariposa Co., Calif. 16. IV. 61, ca. 1200’, 
J. W. Tilden. 

Callophrys viridis Edw. Twin Peaks, San Francisco, Calif. 16. III. 57, J. W. 
Tilden. 

Callophrys affinis Edw. Dry Canyon, Salt Lake County, Utah 14.VII.49, J. C. 
Downey. 


Photography by Jean Norton 


290 


TILDEN 


J. Res. Lepid. 


tipped and usually contrasting, particularly on the hind wings. Occasion- 
al females in a long series are brown, very seldom fulvous, and nor- 
mally females cannot be separated from males on color alone. The 
outlines of the wings are more quadrate than in other members of 
the subgenus; the crenations of the hind wings are discernible but 
small. The forewing below has the gray limited to the anal margin, 
not invading the disc. The green below is dark and frequently bluish, 
quite distinctive. The antennae are pale to whitish above, seemingly an 
unique character. The range in California is rather narrowly limited 
to the coast, from Santa Cruz County northward. Its range beyond 
California needs to be clarified. It flies very early in the year, late 
February to early April, and seldom leaves the vicinity of Eriogonum 
(usually E. latifolium Sm.) except to visit nearby flowers. It is partial 
to flowers of Umbelliferae. Its flight is low and easily overlooked. In 
spite of reports to the contrary, viridis seems to be the only species of 
the subgenus found in the immediate environs of San Francisco, and 
is also common in similar habitats in Marin County. 


Gallop hrys af finis ( Edw. ) 

Proc. Acad. Nat. Sci. Phila., 223:1862 
A fulvous to bright rufous insect, with dark terminal line and 
usually dark contrasting stigma, the fringes pale-tipped to white, 
usually very contrasting. Below, uniformly pale yellow-green (nom- 
inate af finis), the macular band obsolete or represented by one or two 
minute macules only. This describes nominate material from Utah. 
North and west, specimens associated with this species are less fulvous 
above, more bluish-green below. W ashingtonia Clench (1944) is based 
upon such specimens from Brewster, Washington. 


Callophrys sheridanii ( Edw. ) 

Eield and Eorest 3:48:1877 

Whether this name is to be attributed to Edwards or to Carpenter 
may be a matter of opinion, A rather short badly written article with 
several misspellings and typographical errors, by Carpenter, states that 
Edwards is describing the species. It reads: 'Theda sheridonii (sic), 
new species, is named in honor of Lieut. Gen. P. H. Sheridan, U. S. 
Army, by W. H. Edwards, Coalberg, West Virginia, at the request 
of W. L. Carpenter, U. S. Army. Size and form ...” Inasmuch 
as it is expressly stated that Edwards is writing the description and 
that it is at the request of Carpenter, the position is taken here that 
Edwards is the author of the name. However, the alternate opinion 
has also been expressed. In any case the present spelling is an emenda- 
tion of an evident lapsus calami. 


(4) s 28 i -30 o , 1963 


NORTH AMERICAN SPECIE^ 


291 



Fig. 2. Callophrys. Upper side to left; lower side to right, 

Callophrys sheridanii Edw. Flagstaff Mt., Boulder Co., Colo. 17.IV.52, Don 
Eff. 

Callophrys apama Edw, Strayhorse, Greenlee County, Ariz., 7.VII.58, 7800 
ft., J. W. Tilden. 

Callophrys comstocki Henne Providence Mts., San Bernardino Co., Calif. 2. IV. 
50, Ray Hulbirt. 

Callophrys lemberti Tilden, n. sp. West above Tioga Pass, Yosemite National 
Park, Calif. 8.VII.58 ca. 10500 ft., O. Shields. 

Photography by Jean Norton 


292 


TILDEN 


/. Res. Lepid. 


Sheridanii is the most divergent species in an otherwise closely re- 
lated group. It is dark gray above in both sexes, with a black terminal 
line and white-tipped contrasting fringes. The stigma of the male 
is small, usually concolorous and not immediately evident. Below, 
the green is very dark and mixed almost evenly with black scales. The 
forewing is more acute than in the other species and the secondaries 
appear smaller and more evenly rounded. The facial tuft is unusually 
dense, erect and black, and most specimens show eighteen annuli in 
the antennae. The macular band is only slightly arcuate and is not 
usually broken into discrete macules. It is white mesially, faced on both 
edges with black. The name neoperplexa B. & McD. (Contrib. 5:671;- 
1923 ) was applied to specimens from Utah in which the macular 
band tends to become reduced or obsolete. The western range of 
neoperplexa remains to be established, but it appears to extend into 
eastern Oregon and eastern Washington. 

Callopbrys cornstocki Henne 
Bull. So. Calif. Acad. Sci. 39:71:1940 

This species was described from the desert region of San Bernardino 
County, Calif. (Providence Mountains) and specimens are relatively 
scarce in collections. The precarious climate results in good populations 
of adults only in favorable years. Comstocki has been considered either 
a distinct species or a subspecies of dumetornm. Examination of the 
short series available to me for study, while not conclusive, indicates 
that co7nstocki is separable from other named segregates of the sub- 
genus by characters at least equal to those defining most of the species. 
It is a gray insect in both sexes, the terminal line dark, the fringes 
white tipped and contrasting at least on the secondaries. The green 
below is dark, the veins slightly darker than the background color. 
The stigma is small, scarcely discernible and the facial tuft is dense, 
coarse, black and proclinate, almost concealing the underscaling. The 
vein tips are not dark on the hind wing below, but the terminal line 
there is black, nearly complete and inwardly bordered narrowly with 
white. The forewing is deeply invaded by gray, a character (almost 
the only one) it shares wiuh dimietoriim. The macular band is com- 
plete but of discrete macules, inwardly black, outwardly white, and 
the spot in cell Cu 2 is displaced outwardly. 

Callopbrys lemberti Tilden, n. sp. 

A mouse-gray smoothly scaled species with pale or concolorous 
stigma and contrasting white-tipped fringes. Sexes similar, or females 
dull brown. Green of lower surfaces pale, the scaling thin, the ground 
color showing through. Macular band complete to obsolete, unusually 
narrow, of discrete macules. 

Holotype male: Costa of forewing 13 mm.; costa upcurved to basal 


(4):28r-300, 1963 


NORTH AMERICAN SPECIES 


293 



Fig. 3. Map showing distribution of CaUophrys apama (°) and C. sheri- 
danii ( • ) . 


294 


TILDEN 


/. Res. Lepid. 


third of cell, then straight to end of R2 + 3, curved down thence to 
apex; outer margin of forewing curved to M3, thence nearly straight 
to 2nd A except for a slight indentation between Cui and Cus; tornus 
diagonal, anal margin nearly straight. Hindwing: Rs=2W A; Mi = 
Cui, these the longest veins in this wing; two slight marginal crenations 
between Cui and Cu2 and between Cuo and 2nd A; white annuli of 
antennae 16 as seen from lateral view; palpi dark above, grizzled 
black and white on sides and below, rather more blunt than usual 
in the subgenus but of no significance, since other specimens of the 
type series have palpi more pointed than in the holotype; facial area 
with gray, slightly proclinate hairs, the vestiture more dense laterally 
and with few irridescent green subscales showing through; body dark 
above, pale below; the legs annulated dark gray and white. 

Upper wing surfaces gray (nearly mouse gray), the veins very 
slightly darker but not greatly contrasting; stigma nearly concolorous 
with wing; fringes of forewing concolorous at base, white at tips; 
fringes of hind wing concolorous gray mixed with golden brown scales 
at base, snow-white and contrasting at tips; tornal table downturned 
and dark. 

Lower wing surfaces with smooth, pale, slightly yellowish green 
(nearly apple green) overscaling, which is thin, the gray ground 
color showing through between the individual scales, costa of fore- 
wing pale brown, moderately constrasting; anal area of forewing 
gray from margin to Cu2, the gray not invading the disc; forewing 
with a slight suggestion of ' a macular band with macules in cells 
M3, Cui and Cu2; fringes as on upper surface. 

Secondaries wdth green overscaling over entire surfaces; macular 
band nearly complete but narrow, the macules narrowly black in- 
wardly, outwardly and more widely (about Vd) white; the macule in 
cell ^rd A is a short dash; that in cell 2nd A directed diagonally toward 
wing base; that in cell Cui. also diagonal but displaced towards wing 
margin by about one-half its own length; macule in cell Cui lacking 
on right wing, indicated by four or five white scales on left wing; 
that in cell M3 very faint, dull white and narrow; that in cell M2 
faintly indicated by a lack of green overscaling only; no macule in 
cell Mi; macule in cell Rs small but distinct; fringes nearly as 
on upper surface except for clusters of dark scales at vein tips, sug- 
gesting incomplete checkering; the tornal tab centrally black, narrow- 
ly faced on each side with white hairs; hairs of vannal margin gray 
at end of macular band, thence nearly white to base. 

Allotype female: Forewing costa 13.5; wing shape and proportions 
essentially as in holotype male; white annuli of antennae 16 (an 
incomplete 17th on base of club); facial area with hairs sparse (as 
is frequent in female Callophrys) ; Body and legs as in holotype male. 

Upper surfaces dull gray-brown, darker and very slightly more 


(4):28 i -3 oo , 1963 


NORTH AMERICAN SPECIES 


295 



Fig. 4. Map showing distribution of Callophrys viridis (1), lemberti (2), 
and comstocki ( 3 ). 


296 


TILDEN 


/. Res. Lepid. 


brownish than holotype male. Costa above with warm brown scales, 
some of which invade the wing nearly to vein Sc; fringes of fore- 
wings dark gray at base, white at tips, of hind wings brownish at base 
except for dark gray patches at vein tips, the fringe tips white; tab 
down-turned and dark, the fringes in the tornal area invaded by 
gray, the tips less contrasting. 

Green overscaling of lower surfaces as in the male; forewing 
costa warm brown (nearly cinnamon); macular band indicated by 
two indistinct macules, in cells M3 and Cu,; fringes light gray at 
base, with darkenings at vein tips; fringe tips white; gray anal margin 
free of green overscaling not quite to Cus; macular band of hind 
wings very narrow, the spots small and white with the inward black 
scaling scarcely more than suggested, the band slightly bowed out 
but the macules not noticeably displaced; macules in cells M2 and M3 
obsolete; fringes pale gray, distinctly dark at vein ends, appearing 
checkered; fringe tips white; a faint subterminal row of paler green 
scales before the fringes; tab and vannal margin as in holotype male. 

Type material: Holotype male. West above Tioga Pass, Yosemite 
National Park, Calif., 9:VII.62, leg. Oakley Shields; allotype female, 
same locality, 10.VII.58, leg. Oakley Shields; seven designated para- 
types as follows: 1 male 8.VII.58 (Shields); 2 males, + 1 $ 19. VII. 52 
(Tilden); I male 25.VI.6l, -j- 1 9 (Dirks); 1 male, 25.VI.62 
(Dirks): All paratypes from same locality. Type locality: West above 
Tioga Pass, about 1 mile, where the Gaylor Lakes Trail reaches its 
highest point before dropping down to Gaylor Basin, thence southerly 
along the ridge to rock oucrops, about two to three hundred yards. 

Type material distributed as follows: Holotype male and allotype 
female in the collections of the California Academy of Sciences; one 
male paratype in the collection of Oakley Shields, La Mesa, Calif., 
one male paratype in the collection of the Los Angeles County 
Museum; one female paratype in the collection of the Carnegie 
Museum, Pittsburgh, Penn.; one male paratype in the collections 
of the National Museum, Washington, D. C. The males are retained 
by the author because they have been dissected to examine the geni- 
talia. Certain other specimens from the type locality are at hand but 
are in too poor condition to form paratypes. 

Variation in the type series: In flown specimens the green appears 
very slightly darker; the macular band is never more complete or 
conspicuous than in the types. In three paratypes the macules of the 
band are smaller and in one specimen the band is very faint. One 
male paratype has the stigma very pale; in one other the stigma is 
slightly darker than the ground color. 

Recognition characters for Callophrys lemherti, n. sp. are the smooth 
gray upper surfaces, nearly similar in both sexes; the very contrasting 
white-tipped fringes, particularly on the hind wings; the very narrow 


NORTH AMERICAN SPECIES 


297 


I (4) ;28i-3oo, 


1963 



Fig. 5. Map showing distribution of Cdlophrys af finis. 


298 


TILDEN 


J. Res. Lepid. 


but frequently nearly complete macular band; the sub-checkered 
fringes of the hind wings below. It may be separated at a glance from 
dumetorum by the smooth gray upper surfaces and white fringes, 
and by the restriction of the gray of the lower surfaces of the 
hind wing to the anal margin. From viridis it may be separated by 
smaller size, more trigonate forewing and the pale green scaling below. 
From comstocki is may be known by the lack of contrast between 
ground color and the wing veins, especially on the lower surfaces of 
the hind wings, and by the more regular and narrower macular 
band and the contrasting brown coloration of the forewing costa. It 
bears no confusing resemblance to the remaining species. Specimens 
of lemherti, n. sp., collected long ago, tend to fade, the females parti- 
cularly showing a sordid gray-brown above. The tendency of Callo- 
phrys specimens to fade is general. 

Specimens from other localities than the type locality referred 
to lemberti, n. sp., are: I male, 1 female, west slope Mt. Dana, Yose- 
mite National Park (Tilden); 1 male, I female, Warren Creek, Mono 
County, Calif. (Shields); 1 female. Mammoth Crest, Mono Co., Calif. 
( J. Powell ) ; 2 males 1 female. Chipmunk Flat, Tuolumne County, 
Calif. (J. Powell); 2 males 1 female, the knobs just north of Sonora 
Pass, Stanislaus County, Calif. ( Shields ) ; 1 male 6 females, Ebbetts 
Pass, Alpine County, Calif. (J. Powell); 1 male I female, Leviathan 
Peak, Alpine County, Calif. (J. Powell); 1 male. Echo Lakes Area, 
Eldorado County, Calif. (Dirks); another male from Echo Lakes, 
without collector label; 1 female, Mt. Tallac, Eldorado County, Calif. 
(F. X. Williams, 1909); 1 male. Tamarack Lake, Eldorado County, 
Calif, (no collector label); 1 male. Crater Lake National Park, Kla- 
math County, Oregon (D. Huntzinger); 1 male, west slope Mt. 
Thielson, Klamath County, Oregon (Shields), 26 specimens in all 
extending from the Central Sierra Nevada to nearly central Oregon. 
In addition there are several apparently conspecific specimens too 
worn to use for reference. 

The species is named in honor of an early collector in the Tuolumne 
Meadows and Tioga Pass sections of what is now Yosemite National 
Park. 

Thanks are due the following individuals and institutions for the 
loan of material: Ernst Dornfeld, Corvallis, Oregon; David Huntzinger, 
Mt. Timpanogos Cave National Monument, Utah; C. Don MacNeill, 
California Academy of Sciences, San Francisco, Calif.; E. J. New- 
comer, Yakima, Washington; Jerry Powell, California Insect Survey, 
University of California, Berkeley, Calif.; Oakley Shields, La Mesa, 
Calif., and Fred Thorne, El Cajon, Calif. 

Special thanks are due Paddy McHenry, Burbank, Calif., for pro- 
viding certain references not otherwise available. 


C4)s2Si-30O, 1963 


NORTH AMERICAN SPECIES 



Fig. 6. Map showing distribution of Callophrys dumetomm. 


300 


TILDEM 


/. Res. Lepid. 


SUMMARY 

The species of Callophrys Billberg, s. str., are compared by use of 
certain minor characters mostly previously unused. An unrecognized 
species, C. lemherti, n. sp., is described. Specimens other than those 
of the type series are referred to this new name and the range as 
now known is given, being from the central Sierra Nevada of Cali- 
fornia to Mt. Thielson, Oregon. 


REFERENCES 

BARNES, W., & J. McDUNNOUGH 1923. Contrih. Nat. Hist. Lepid. North 
America. 5:64. 

BILLBERG, C. J. 1820. Enumeratio insectorum in Museo Billberg, p. 80. 

BOISDUVAL, JEAN A. 1852. Lepidopteres de California. Ann. Soc. Ent. 
France, (2) 10:291. 

CLENCH, H. 1944. Notes on lycaenid butterflies, a. The genus Callophrys 
in North America. Bull. Mus. Comp. Uoo. 94:217-229. 

CLENCH, H., in EHRLICH, P. R., & ANNE H. EHRLICH. 1961. How to 
know the Butterflies, 200-211. 

EHRLICH, P. R. & H. CLENCH. I960. A new subgenus and species of 
Callophrys (s. 1.) from the southwestern United States. Ent. News 
71:137-141. 

EDWARDS, W. H. 1862. Proc. Acad. Nat. Sci., Philadelphia, p. 223. 

. (in CARPENTER, W. L.) 1877. Field & Forest, 3:48. 

. 1882. Papilio 2:137. 

HENNE, C. 1940. Bull. So. Calif. Acad. Sci., 39:71. 

LINNEAEUS, C. 1758. Systema Naturae, 10th Ed., 1.483. 

OBERTHUR, C. 1913. Etudes Lep. Comp., Ease. IX (Partie Ire) PI. CCXXX- 
VI, fig- 1926. 

SCUDDER, S. 1875. Proc. Am. Acad. Arts & Sci., Boston, 10:132, no. 202. 

ZIEGLER, J. B. i 960 . Preliminary redefinition of North American Hair- 
streak genera. J. Lep. Soc., 14:19-23. 


THE FOUNDATION exists as a charitable fund to provide per- 
manent security to the Journal. Funds taken in are placed in a place 
or places where they will obtain reasonable earning power together 
with safety. Regular membership in the Foundation is 10.00 per year 
(coinciding with the Journal volume); other memberships are avail- 
able. See the special form later in this issue. The Foundation will also 
maintain collections suitable for geographical or biological study; gifts 
are welcomed from anyone. No public exhibits are planned. The 
Foundation welcomes help in its cause; it does not feel competitive with 
any other group but rather complementary. Arrangements for pay- 
ment in foreign currencies are possible; write to the editor for in- 
formation. 


YOU ARE CORDIALLY INVITED 

TO BECOME A MEMBER OF THE LEPIDOPTERA FOUNDATION, WHICH IS 
DEDICATED TO THE ADVANCEMENT OF KNOWLEDGE OF MOTHS AND 
BUTTERFLIES THROUGHOUT THE WORLD, THROUGH THE PUBLICATION 
OF THE JOURNAL OF RESEARCH ON THE LEPIDOPTERA. 


Membership includes subscription to the JOURNAL and a donation of the balance 
of the subscription to a fund which will be used for a permanent endowment. 


This Page Can Be Cut Out Without Defacing The Issue. 


THE LEPIDOPTERA FOUNDATION 

1140 West Orange Grove Ave., Arcadia, California, U.S.A. 


Application for membership: 

Name 

Address 


hereby applies for 

membership in the Foundation. 


CLASSES OF 
MEMBERSHIP 


Regular 

Family 

Contributing 

Subscribing 

Sponsor 

Life^ 


$ 10 year 
15 year 
25 year 
50 year 
100 year 
250 for Efe 


*Life membership includes the Journal for your life; should any future change occur such 

that the Journal would cease publication, the full amount of the Life membership will 

be returned to you during your lifetime. 

The Foundation will receive bequests or gifts in the form of funds, property 
or colleaions. 

301 


NOTICE 


MEETING: Western Section, Lepidopterists Society- 

Place: Santa Barbara Museum of Natural History, 
Santa Barbara, Calif, 

WHEN: August 24-25, 1963. 

Papers: Over ten papers and demonstrations have 

already been received; there is time 
for more to be received. Send to the 
Editor of this Journal. 

IN ADDITION: THE LEPIDOPTERA FOUNDATION, 
its aims and purposes will be 
discussed. 


302 




FOR COLLECTING WITH LIGHTS 
12 1/2 LB POWER SOURCE 


Weighs only 12 1/2 lbs„ 

2 cycle, 1 3/4 H*P. 

Direct Drive 

Ea sy starting 

Air cooled 

Made in U.S.A. 

Guaranteed 90 days 

Uses standard outboard fuel 

Roller and ball bearings 

Completely self contained 

World wide service available 

#5830 Tiny Tiger generator, 

115 volt AC, 12 volt DC, 30 0 watts $99.50 

#5831 Carrying case 8.9 5 

Prices listed are f.o.b. Santa Monica, Calif. 
California residents please add 4% sales tax. 

B\0 METAL ASSOCIATES 

Box 61 

Santa Monica, California 




THE JOUI^NJAL OF KESIAR^OHJ 
ONJ THE LEFIJOOFTERA 


Volume 1 Number 4 May, 1963 

IN THIS ISSUE 

Techniques in the study of population structure in 
Philotes sonorensis 

Rudolf H, T. Mattoni and Marvin S. B. Seiger 237 

Early stages of a southern California Geometrid moth, 

Drepamdatrix hulsti huhti (Dyar) John Adams Comstock 245 

The effectiveness of different isothiocyanates on attracting larvae 
of Eteris rapae 

William Hovanitz, Vincent C. S. Chang and Gerald Honch 249 

The origin of a sympatric species in Colias through the aid of 

natural hybridization William Hovanitz 261 

A method for breeding Pieris napi and Pieris hryoniae 

Bjorn Petersen 275 

An analysis of the North American species of the genus 

Callophrys J. W. Tilden 281 

THE LEPIDOPTERA FOUNDATION 301 

Notice 302 

BIOGRAPHICAL SKETCHES 

A number of these have been received but we need more. The space 
where these could go is lost at the end of papers unless authors send in their 
photos and sketches. We hope to start this feature in volume 2. 




Established in 1 962 


Edited by WILLIAM HOVANITZ 


Volume 2 


1963 


Published at 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 









Volume 2 


Number 1 


July, 1963 


YOSEMITE BUTTERFLIES: An ecological survey of the 
butterflies of the Yosemite sector of the Sierra Nevada, 
California ..... John S. Garth and J. W. Tilden 1 

Volume 2 Number 2 September, 1963 

Quantitative analysis of certain wing and genitalia characters 
of Pieris in western North America 

Vincent C.S. Chang 97 

Biographical sketch — Vincent C.S. Chang 126 

Genetic and environmental variation in Pieris brassicae 

Brian O.C. Gardiner 127 

Type localities of the Megathymidae H. A. Freeman 137 

Biographical sketch — H. A. Freeman 142 

The distribution of an endemic butterfly Lycaena hermes 

Fred Thorne 143 

Callophrys (Lycaenidae) from the Northwest 

Harry K. Clench 151 

Euphyes dukesi Bryant Mather I 6 I 

The complete life history of Staphylus hayhursti 

John Richard Heitzman 170 


Volume 2 


Number 3 


November, 1963 


Affinities and distribution of Antillean Ithomiidae 

Richard M. Fox 173 

Ovipositional preference tests with Pieris 

William Hovanitz and Vincent C.S. Chang 185 

Notes on the early stages of Drepanulatrix monicaria 

(Guenee) ( Geometridae ) John Adams Comstock 201 

Biographical sketch — John Adams Comstock 204 

The origin of a sympatric species in Colias througli the aid 

of natural hybridization William Hovanitz 199 

Biographical sketch — James Wilson Tilden 224 

A rubber stamp method for producing specimen labels 

Phillip A. Adams 225 

Generic or subgeneric names closely related to Argynnis 

Paddy McHenry 229 

THE LEPIDOPTERA FOUNDATION 240 

Volume 2 Number 4 December, 1963 

Evidence for lack of territoriality in two species of Hamadryas 

Gary N. Ross 241 

A synopsis of the west Indian Lycaenidae, with remarks on 

their zoogeography Harry K. Clench 247 

The synonymy, variability and biology of Lycaena nivalis 

E. J. Newcomer 271 

Comparison of the selective effect of two mustard oils and 
their glucosides to Pieris larvae 

W. Hovanitz and V.C.S. Chang 281 

The Annaphila astrologa complex, with descriptions of three 

new species Frank P. Sala 289 

Old master painting Clark, Nelson of London, Ltd. 302 

Dates of publication of volumes 1 and 2 302 

Collecting of Annaphila spila with notes on the "crimson- 

winged” group of the genus J. S. Buckett 303 



Volume 2 


Number 1 


]ul^, 1963 




THE JOUKNJAL @F RISEAKCHJ ©H 
THE LEPIJDOFTEKA 

a quarterly published at 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
edited by: WILLIAM HOVANITZ 

THE PURPOSE OF THE JOURNAL is to combine in one source 
the work in this field for the aid of students of this group of insects 
in a way not at present available. THE JOURNAL will attempt to 
publish primarily only critical and complete papers of an analytical 
nature, though there will be a limited section devoted to shorter papers 
and notes. QUALITY WORK on any aspects of research on the 
Lepidoptera is invited. Analytical and well illustrated works are pre- 
ferred, with a minimum of long description. 

AUTHORS ARE REQUESTED to refer to the journal as an 
example of the form to be used in preparing their manuscripts. Illu- 
strations should be of the best quality black and white, or line draw- 
ings and should be pre-arranged by the author to fit a reduced size 
of 4” X 6V2.” Footnotes should be avoided; bibliography should be as 
indicated. Tables should be set-up for page size as indicated. Manuscripts 
in good form and requiring little work by the editor will naturally 
appear first. Authors, who wish drawings made for them, may submit 
rough sketches and will be billed for the cost, which will be very 
negligible. Work to be done on res^rch grants should so specify. When 
possible, tabular matter should be typed on good paper with a carbon 
ribbon in a form suitable for a one-third reduction and in a size to 
fit 4” X 61/2.” 

THE JOURNAL is not a suitable place for continued changes 
of nomenclature; unless the author is himself analytically studying a 
group from its biological point of view and finds a change necessary, 
the editor must ask authors to refrain from any changes from the 
McDunnough Check List unless superseded by a monograph published 
since that date. Popular books are not to be considered as giving scien- 
tific credence to any name. It is rare that name changes need be made 
and preference is given to old names unless in the editor s opinion 
sufficient evidence is given to warrant such change. 

SUBSCRIPTIpNS should be sent to the above address. 

RATES are: $8.00 per volume, personal subscription (but see below) 
$12.00 per volume, institutional subscription. 

The personal subscription rate is included in the membership to the 
Lepidoptera Foundation indicated below. SPECIAL SERVICE TO 
FOREIGN ADDRESSES: THE PURNAL will be mailed air mail 
or registered at cost to the subscriber, if so desired. 


2(l):l-96 


Journal of Research on the hepidoptera 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
Copyright 1963 


YOSEMITE BUTTERFLIES 


An Ecological Survey of the Butterflies of the 
Yosemite Sector of the Sierra Nevada, California 


JOHN S. GARTH 

Allan Hancock Foundation 
University of Southern Calif orna 
Los Angeles, California 

and 

J. W. TILDEN 

San Jose State College 
San Jose, California 


July, 1963 


2 


GARTH AND TILDEN 


/. Res. Lepid. 


FOREWORD 

Organized interpretation of the chief features of National Parks had its 
beginning in Yosemite National Park in 1920. Suitable guide leaflets and 
bulletins found a useful place in the educational program. At first printed 
material covered wide fields of nature study. But with more visitors have 
come many with specialized backgrounds and they seek help. Whereas, the 
first bulletins attempted coverage of all insects found, we now come to one 
dealing only with butterflies. The senior author graduated from the Yosemite 
School of Field Natural History with the Class of 1933 and made his first 
contribution to the subject by publishing a list of the Butterflies of the 
Boundary Hill Research Reserve (Bull. Southern California Academy of 
Sciences, 33, Sept.-Dee., 1934, pp, 13T135). He widened his studies to the 
Butterflies of Yosemite National Park in the fifties and herewith is a paper that 
emphasizes where each butterfly is found and what plant its larvae feed upon. 
Colored plates give the amateur a chance to identify species found and the 
text places zonal locations and ecology. Other national parks that have 
profited by the senior author’s field work are Grand Canyon, Glacier, and 
Grand Teton. Crater Lake and Lassen have profited similarly from the field 
work of the junior author.- — Harold C. Bryant. 


The original costs of this publication were underwritten by the 
Lepidoptera Foundation and the Journal of Research on the Lepidoptera 
and the separate issue is made available as a public service to visitors of 
the Yosemite National Park and adjacent areas. 


William Hovanitz, Founder, The Lepidoptera Foundation 
Editor, The Journal of Research on the Lepidoptera 


(c) Copyright 1963 

Additional copies may be obtained by writing to THE LEPIDOPTERA 
FOUNDATION, 1140 W. Orange Grove Avenue, Arcadia, California. 


ALLAN HANCOCK FOUNDATION CONTRIBUTION Number 248. 


2(i):i-$6, 1963 


YOSEMITE BUTTERFLIES 


3 


CONTENTS 

Foreword 2 

Contents 3 

Historical Note 3 

Introduaion 5 

The Lepidopterist’s Yosemite 6 

The Surrounding National Forests 7 

Yosemite Foot-Trails 8 

Life-zones 9 

Butterflies as Life-zone Indicators 9 

Biotic Provinces .......................................................................13 

Plant Communities 14 

Food Plants ...........................................16 

Some Plants on Which Yosemite Butterflies Feed as Larvae ..........17 

The Geological Background ..........18 

Acknowledgments ..............19 

Names Used in This Paper ..19 

Account of Species of the Yosemite Region 22 

Literature Cited ........................68 

Yosemite Butterfly Records ............................70 

Appendix 1. Food Plant Records ....................................................71 

Three Anomalous Situations ............................................................................77 

Appendix 11. Yosemite Records ......................................................................78 

Api^ndix III. Localities ..................................................................................92 

Lepidoptera Foundation ....................................................................................96 


HISTORICAL NOTE 

The authors are indebted to Mrs. Mary V. Hood, Park Historian, 
Yosemite National Park, for calling their attention to an early record 
of butterfly collecting by John Muir, as recorded by W. F. Bade in 
Life and Letters of John Muir, 1924: 

Muir to Mrs. Ezra S. Carr, August 13, 1871. "I suppose you have seen 
Mr. King, who kindly carried some [butter]flies for Mr. Edwards ... I collert- 
ed most of them upon Mount Hoffman, but was so busy in assisting Reilly 
that I could not do much in butterflies.” (Op. cit., Vol. I, p. 292). 

Henry Edwards to John Muir, August 25, 1871. "In the small box that 
you sent me [through Mrs. Carr] are four species new to my collection and two 
of these ate new to science . . . All the specimens are rare, and are different 
from those found in the Valley. The two new species are the bright crimson 
copper one from Cathedral Peak, and one of the small bluish butterflies. There 
is a pair of greenish yellow ones, very rare and interesting. The species was 
described from a pair only which were taken by the Geological Survey at the 
headwaters of the Tuolumne River, and strange to say, no others have turned 
up until you found it now ... It is really very singular that the remove of a 
few miles from the Yosemite should produce species so very different from 
those of the Valley itself, and at the same time so charaaeristic in their forms.” 
(Op. cit., pp. 263-264). 

Of the butterflies mentioned, the bright crimson copper is probably 
The Lustrous Copper, Lycaena cupreus (Edw.), while the greenish yel- 
low ones are undoubtedly Behr’s Sulphur, Colias behrii Edw. It should 
be noted, however, that the author is W. H. Edwards in each case. 
Both species are illustrated on color plate III. 


I^IITBTICS 


r,i i; (I r't 



4 


GARTH AND TILDEN 


/. Ris. Lepid. 



Frontispiece. Boundary Ridge, el. 9,000 ft.. Research Reserve. Western 
White Pine and sparse ground cover of herbs and grasses. Hudsonian life 
zone elements weakly present. Rocky outcrop association. Papilio zelicaon 
flies to the summit, while Euchloe creusa hyantis and E. ausonides coloradensis 
keep to the leeward of the ridge.— -Neva Snell. 


3( i ): i -96, 196} 


YOSEMITE BUTTERFLIES 


5 


INTRODUCTION 


The Yosemite National Park embraces 1,189 square miles of the 
most diversified territory to be found in the Sierra Nevada. From the 
edge of the San Joaquin Valley at near sea level it extends to the crest 
of the Sierran divide, culminating in Mt. Conness, Mt. Dana, and Mt. 
Lyell, each over 13,000 feet in elevation. Between these extremes 
occurs a wide range of climatic conditions, giving rise to a wealth of 
plant and animal life scarcely to be duplicated elsewhere on the North 
American continent. The vertebrate fauna has been thoroughly investi- 
gated by Grinnell and Storer (1924), who recorded 97 species of 
mammals, 231 of birds, and 34 of reptiles and amphibians in "Animal 
Life in the Yosemite," their Yosemite sector extending beyond park 
boundaries to include part of the San Joaquin Valley and of the Mono 
Basin as well. 

The larger host of invertebrates, and of insects in particular, have 
long demanded similar investigation. The butterflies are an ideal group 
with which to begin this study. They are diurnal, or daylight fliers, and 
so lend themselves to easy observation, as distinguished from moths, 
of which most are nocturnal or crepuscular in flight. They are also for 
the most part conspicuous, and therefore likely to attract the attention 
of even the casual park visitor. Again, their number as species found 
in a given area roughly approximates the number of bird species, if 
only those actually residing (i.e., nesting) in that region be enumer- 
ated. Thus the butterflies constitute a natural group of insects small 
enough to be encompassed by the amateur naturalist in a season, yet 
sufficiently diversified to entice the professional entomologist to return 
year after year to resolve the problems of habit and habitat that they 
present. 

Like the many other natural attractions of our National Parks, but- 
terflies enjoy the protection of far-sighted legislation and are held in 
trust for future generations as part of America s outdoor heritage. 
Their unauthorized removal is contrary to park regulations; however, 
the advanced researcher who can establish his connection with a public 
museum or other scientific institution, such as a university, may qualify 
for one of two types of collecting permits, and should consult the Park 
Naturalist, with offices in the Yosemite Museum, regarding his eligi- 
bility for this consideration. 


6 


GARTH AND TILDEN 


/. Res. Lepid. 


THE LEPIDOPTERIST’S YOSEMITE 

The territory now included in Yosemite National Park has been 
known to butterfly enthusiasts since the days of the gold rush of ’49. 
The pioneer entomologist, Pierre Lorquin ( for whom Lorquin’s 
Admiral is named), followed the mother lode to Placerville, and from 
thence crossed the Sierra. His exact route is not known, but if he 
crossed by Tioga Pass, el. 9,941 ft., it is quite possible that the "Great 
Salt Lake" above which he discovered the Irene Fritillary (Argynnis 
atlantis trene) was Mono Lake, and not the Great Salt Lake of Utah. 
Indeed, had type localities been designated as carefully then as now, 
we might find that many of our California butterflies were first ob- 
served by a naturalist somewhere in the Yosemite region. The hermit 
Lembert (whose name is memorialized in Lembert’s Dome) for many 
years supplied the entomological world with specimens of Behr’s 
Sulphur {Colias hehrii) from his hideaway in Tuolumne Meadows, 
and it was not until the Tioga Road opened to tourist travel in 1915 
that his secret source of this rarity was discovered. 

The entire northern section of the park and much of the south- 
eastern part is still traversable by foot or horseback only, and is virgin 
territory for the lepidopterist. For many years following its establish- 
ment in 1925 the Yosemite Field School (short title for Yosemite 
School of Field Natural History) conducted annual summer excursions 
into this hinterland under the capable leadership of former Park 
Naturalist C. A. "Bert" Harwell. Begun as a six-day circuit of the 
established Hi-Sierra Camps, this "back-country trip” developed into 
an independent pack caravan that lost itself among the glaciers for a 
ten-day to two-week period and seldom returned without a contribution 
to the natural history of the park. During the 1933 to 1937 period the 
Field School included among its members a number of students of 
insects whose training and experience made it possible to observe and 
record much valuable information concerning the habits of rarely en- 
countered species, particularly in connection with the annual surveys of 
Boundary Hill Research Reserve, instigated in 1933 by Joseph S. Dixon, 
field biologist, National Park Service. Sections on butterflies in the 
Research Reserve Reports were written in 1933 by John S. Garth and 
Fred C. Ziesenhenne, in 1934 by Janet Mabry and Edmund D. God- 
win, in 1935 by Theodore H. Eaton, Jr., and in 1937 by Robert 
Godwin, Frances Cramer, and Verlin Baysinger. The 1936 report, by 
R. Paul Allen and Carsten Aherns, noted succinctly: "Rain for six 
days, no butterflies.” These reports are on file at the Yosemite Museum, 
where they constitute an important supplement to the published report 
of Garth ( 1935a) . 

It was as a resurvey after an absence of 23 years from Yosemite that 
the senior author undertook the present assignment at the request of 
Chief Park Naturalist Douglass H. Hubbard, the intervening years 


a(i):i~96, 1963 


YOSEMITE BUTTERFLIES 


7 


having been enriched with similar experiences at Grand Canyon, 
Glacier, and Grand Teton national parks. Desirous of extending the 
earlier coverage of the park, he began his observations in 1956 in the 
vicinity of Mather, including the lower Tuolumne, Hetch-Hetchy, and 
Aspen valleys. In the same year the junior author, whose Yosemite 
experience also dates from the late 1920’s, began under park auspices 
an ecological study of the area adjacent to Tioga Pass, including a 
transect from Pilot Peak to Mt. Dana (Tilden, 1959). Since both 
investigators were limited to brief visits to the area at critical periods, 
they were fortunate to find in Oakley Shields a summer resident whose 
activities spanned the entire season at Tuolumne Meadows, where his 
father, Allan Shields, was a seasonal naturalist. By combining records 
over the succeeding 7-year period it has been possible to present a fairly 
consistent picture of the situation in the park’s high country, its western 
approaches, and of the Mono Basin to the east, although the coverage 
of some areas is less complete than might have been desired. 

THE SURROUNDING NATIONAL FORESTS 

The visitor driving to and from Yosemite Valley commonly arrives 
by one of three routes from the west and leaves by another. These are 
California State Highways 41 via Coarsegold and Wawona, 140 via 
Mariposa and El Portal (the All-Year Highway), and 120 via Grove- 
land and Buck Meadows. For many miles he travels through wooded 
foothills of the Sierra and Stanislaus National Forests, where oppor- 
tunities for observing butterflies are equal to those within the National 
Park, while collecting is unrestricted. If visiting Tuolumne Meadows, 
he may wish to leave the Park by its eastern entrance, traveling 
through Inyo National Forest from Tioga Pass to Lee Vining, there 
joining U. S. Highway 395. Should he seek an area comparable to the 
Park s high country he will find it at Saddlebag Lake, accessible by 
road from Tioga Lake, and adjacent to the Hoover Wildlife Preserve 
and the Hall Primitive Area. Other east-slope mountain lakes accessible 
from Highway 395 are Lundy Lake, the Virginia Lakes, and the June 
Lake circuit. 

During the years 1950-1958 the junior writer spent considerable 
time in the Mono Lake region, obtaining many records later verified 
and extended by Oakley Shields. In 1958 the senior writer established 
temporary residence at June Lake and from this base made one-day 
trips to Gem Lake, Saddlebag Lake, and the Virginia Lakes basin. In 
1957 and again in 1959 Oakley Shields crossed the Sierran Divide at 
Donohue Pass and proceeded southeastward via Agnew Pass to Agnew 
Meadows. In the spring of 1961 particular effort was made by Park 
Naturalist Keith Trexler to explore the western foothills, where the 
season begins from six weeks to two months earlier than in Yosemite 
Valley. The results of these extra-limital investigations were to add a 


8 


GARTH AND TILDEN 


/. Res. Ltpid. 


number of species that rarely, if ever, occur within Park boundaries, 
but which deserve consideration in any account of the butterflies of 
the Yosemite region, which includes the encircling National Forest 
lands as well. It is the authors’ hope that the inclusion of these locali- 
ties outside the Park will serve to entice the butterfly enthusiast away 
from the few crowded centers and into the less frequented but no less 
rewarding byways. 


YOSEMITE FOOT-TRAILS 

While some of the most spectacular butterflies, including the Leto 
Fritillary (Argynnis cybele leto) and the California Sister {Limenitis 
bredowii calif ornica) , fly within the confines of Yosemite Valley, the 
lepidopterist seeking the rarities for which the Park is famous must 
be willing to do some strenuous hiking. The valley walls rise per- 
pendicularly three thousand feet and more, but once their summit is 
gained a large expanse of comparatively level country is accessible by 
means of foot-trails that wind through alpine meadows fragrant with 
wildf lowers. Over 700 miles of such trails traverse the territory within 
Park boundaries, making the Yosemite a veritable entomologist’s para- 
dise. 

Leaving the north side of the valley floor by the Yosemite Falls 
trail, a two-hour climb, best accomplished in the early morning, places 
one in a position to select either the Eagle Peak, Yosemite Creek, or 
North Dome trails for a day among the Parnassians ( Parnassius 
clodius baldur and P. phoebis behrii) . Likewise, the ascent to Glacier 
Point by the Ledge or Four-Mile trails allows the entomologist a choice 
of either the Pohono or Glacier Point trails that parallel the south rim 
of the valley and along which the Nivalis Copper {Lycaena nivalis) 
is certain to be encountered. The "high country,’’ reached either via 
Lake Tenaya or Lake Merced, is the habitat of the Ivallda Arctic 
{Oeneis chryxus ivallda), Malcolm’s Checker-Spot {Melitaea damoetas 
malcolmi) , and the Lustrous Copper {Lycaena cupreus) . It was the 
privilege of the senior writer, as a member of the Yosemite Field 
School class of 1933, to cover over 200 miles of trail outside of Yose- 
mite Valley, climbing Mt. Lyell, el. 13,090, Mt. Kuna, el 12,956, and 
Mt. Dana, el. 13,050 feet, on successive days, and in 1958, exactly 25 
years later, to retrace a portion of this itinerary, from Tuolumne 
Meadows to Merced Lake via Vogelsang Pass, with Oakley Shields. 
(And aside from the trail’s being steeper and the butterflies more 
elusive, things were much as he remembered them! ) Generally 
speaking, June is the month for exploring the Yosemite Valley, July 
for the valley’s rim, and August for the glacier country. 


a(i):i-96, 1963 


YOSEMITE BUTTERFLIES 


9 


LIFE ZONES 

The incline from El Portal on the west to the Sierra Crest at the 
eastern park boundary may be subdivided into five regions or life 
zones, each supporting a distinctive flora and fauna. These zones, in 
ascending order, are Upper Sonoran, Transition, Canadian, Hudsonian, 
and Arctic-Alpine. A sixth zone, the Lower Sonoran, occurs a few miles 
west in the San Joaquin Valley. Thus within the greater Yosemite 
region is to be found every life zone recognized in temperate North 
America with the exception of the Subtropical, found only in Florida 
and southern Texas. In many cases the zones merge imperceptibly, as 
when a forest predominantly of Jeffrey Pine and Incense Cedar, typi- 
cally Transition, gives way to one of Red Fir and Quaking Aspen, 
typically Canadian. In others the line of demarcation may be startlingly 
abrupt, as when the south slope of a chaparral-clothed ridge (Upper 
Sonoran) gives way to an open evergreen glade (Transition) on the 
opposite-facing north slope at the same elevation. While the subject 
of life zones has been exhaustively treated elsewhere, it may be said 
that of the factors that determine life zones temperature is the single 
most important, and that temperature is regulated by latitude, altitude, 
direction of prevailing winds, proximity to bodies of water, ascending 
currents of warm air from desert regions, descending drafts of cold 
air from mountain tops or glaciers, character of substrate, and pattern 
of drainage. 

BUTTERFLIES AS LIFE-ZONE INDICATORS 

Plants and animals may be divided into two groups with respect 
to life zones: those that range over a wide territory and those that 
adhere closely to a single zone. The former are called cosmopolites 
because of their unrestricted movement; the latter are called indicators, 
because their presence is considered sufficient to establish the presence 
of the zone. (See GrinneU and Hall, 1919). Thus the Cony {Ochotona) 
of the rock slides is an indicator of Hudsonian Zone; the Sierra Nevada 
Rosy Finch {Leucosticte) of the snow banks is an indicator of Arctic- 
Alpine Zone; and the Chamise {Adenostoma) of the chaparral is an 
indicator of Upper Sonoran Zone. On the assumption that the most 
permanent indicators are those most to be relied upon, plants and 
animals might be arranged, in descending order of reliability, in the 
following order: trees and shrubs, herbs and forbs, mammals, reptiles 
and amphibians, and birds. In attempting to place insects in such a 
scale consideration need be given to their dependence on a particular 
host plant in the larval stage, their normal flight range as adults, and 
their migratory habits. Thus the insect combines the fixity of the plant, 
the mobility of the mammal or reptile, though to a resricted degree, 
and the periodic wanderings of the bird, although less perfectly devel- 
oped. The insects might with some justification be ranked between 


10 


GARTH AND TILDEN 


/. Res. Lepid. 


Figure 1. Profile of the YOSEMITE REGION showing life zones and biotic provinces, together 
with a list of the butterfly species most closely restricted to them. 


CALIFORNIAN PROVINCE 


UPPER SONORAN ZONE 
Coenonympha t. californica 
Cercyonis silvestris 
Argynnis c. inornata 
Euphydryas c. chalcedona 
Euphydryas e. rubicunda 
Melitaea leanira 
Apodemia m. tuolumnensis 
Habrodais grunus 
Strymon auretorum 
Strymon a d e n o s t o m a t i s 
Callophrys d. windi 
Callophrys dumetorum 
Callophrys iroides 
Lycaena a. arota 
Lycaena gorgon 
Philotes speciosa 
Pieris napi venosa 
Epargyreus clarus 
Thorybes pylades 
Heliopetes ericetorum 
Erynnis z. funeralis 

TRANSITION ZONE 
Argynnis c. leto 
Argynnis z. zerene 
Argynnis hydaspe 
Callophrys johnsoni 
Callophrys nelsoni 
Everes amyntula 
Colias o. chrysomelas 
Thorybes diversus 
Hesperia h. yosemite 
Amblyscirtes vialis 


CANADIAN ZONE 
Argynnis irene 
Boloria epithore 
Euphydryas c. sierra 
Melitaea hoffmanni 
Lycaeides a. anna 
Plebejus saepiolus 
Plebejus lupini 
Anthocharis s. Stella 
Pyrgus ruralis 
Erynnis p. lilius 
Hesperia harpalus, blend 

HUDSONIAN ZONE 
Callophrys lemberti 
Lycaena mariposa 
Lycaena nivalis 
Philotes b. battoides 
Plebejus g. podarce 
Glaucopsyche 1. Columbia 
Colias behrii 
Thorybes m. nevada 
Polites s. tecumseh 

ARCTIC- ALPINE ZONE 
Oeneis c. ivallda 
Euphydryas e. nubigena 
Melitaea d. malcolmi 
Lycaena cupreus 
Lycaena p. hyp'ophlaeas 
Plebejus s. comstocki 
Pieris o. calyce 
Parnassius p. behrii 
Hesperia miriamae 


ARTEMESIAN PROVINCE 


CANADIAN - HUDSONIAN 
Cercyonis oeta 
Argynnis z. malcolmi 
Lycaena heteronea 
Lycaena rubidus 
Lycaena editha 
Hemiargus isolus 
Thorybes m. nevada 
Hesperia nevada 

TRANSITION- UPPER SONORAN 
Coenonympha t. mono 
Cercyonis p. ariane 
Danaus g. strigosus 
Argynnis n. apacheana 
Argynnis c. nevadensis 
Euphydryas e. monoensis 
Melitaea acastus 
Limenitis w. nevadae 
Apodemia m. mormo 
Strymon dryope 
Lycaena a. virginiensis 
Everes comyntas 
Leptotes marina 
Philotes b. glaucon 
Pieris beckeri 
Colias p. hagenii 
Hesperia harpalus 


CALIFORNIAN \ ARTEMESIAN 





YOSEMITE BUTTERFLIES 


11 


the plants and the vertebrates, possessing greater reliability as indica- 
tors than the mammals, birds, and reptiles on which much of the eco- 
logical work of the past has been done, their importance in this respect 
having but lately been recognized. 

The first attempts at zonal analysis of California butterflies were 
of necessity incomplete and therefore inconclusive. Based upon the 
record, in some cases, of but a single specimen, and without knowledge, 
in many instances, of the larval food plant, the assignment of the 
species to a given life zone was made without knowledge of its status 
within that zone, whether as resident or vagrant. The pioneer work of 
Comstock and Dammers in the 1930’s made known the life histories of 
most of the butterfly species of the southern half of the state, while 
workers of a younger generation, among whom may be mentioned 
Don MacNeill, John Burns, G. and R. Bohart, Noel McFarland, Paul 
Opler, Tom and John Emmel, and Oakley Shields, have defined the 
range of species and subspecies within the entire state with exactness. 
Thus the life zone normally inhabited by a butterfly of the Yosemite 
region can be stated with more assurance than was possible a quarter- 
century ago. A brief characterization of life zones follows: 

Upper Sonoran Zone: The chaparral or "elfin forest” that clothes 
the foothills from 1,500 to 4,000 feet (occasionally lower) constitutes 
the Upper Sonoran Zone. It is characterized by a great variety of 
shrubby plants, many of which exhibit remarkable adaptations for 
conserving moisture, including anastomosing root systems that check 
soil erosion. Digger Pine, California Buckeye, Poison Oak, and Chamise 
{Adenostoma) are typical species, as is Interior Live Oak of the in- 
cluded Foothill Woodland community. The Chalcedon Checker-Spot 
{Euphydryas c. chalcedona), California Ringlet {Coenonympha tullia 
calif ornica) ^ and Sylvan Satyr {Cercyonis silvestris) seldom stray be- 
yond the confines of this life zone. 

Transition Zone: Between the Austral zones (Lower and Upper 
Sonoran) and the Boreal zones (Canadian, Hudsonian, and Arctic- 
Alpine) lies a broad intermediate area known as Transition Zone. Its 
coniferous forest contains the trees of greatest commercial value: Big 
Tree, Yellow Pine, Jeffrey Pine, Sugar Pine, Incense Cedar, White 
Fir, and Douglas Fir. Azalea, Nuttall Dogwood, and Black Cotton- 
wood fringe its streams; Black Oak and Golden Cup Oak clothe its 
valley floors and mountainsides, respectively. The Leto, Hydaspe, and 
Zerene fritillaries {Argynnis cybele leto, A. hydaspe, and A. z. zerene) 
represent its considerable butterfly population. Transition Zone ex- 
tends from 4,000 (or less in some localities) to 7,000 feet, merging 
imperceptibly into Canadian Zone along its upper border. 

Canadian Zone: Red Fir replaces White Fir in the open forest and 
Quaking Aspen displaces Azalea and Black Cottonwood along the 
water courses. A secondary chaparral, reminiscent of Upper Sonoran 
Zone, but composed of matted Snow Brush {Ceanothm) ^ Chinquapin 


12 


GARTH AND TILDEN 


/. Res. Lepid. 


{Castanopsis) , and Huckleberry Oak {Quercus vaccinifolia) , covers 
the steeper slopes. The Western Banded Elfin {Gallop hrys eryphon) , 
Sierra Checker-Spot {Euphydryas chalcedona sierra)., and Western 
Meadow Fritillary (Boloria epithore) are good butterfly indicators. 
Canadian Zone extends from 7,000 to 9,000 feet (or less in some 
localities), with Lodgepole Pine invading from its upper margin. 

Hudsonian Zone: At about the 8,500 foot level Lodgepole Pine 
appears, to be joined at the 9,000 foot level by Mountain Hemlock to 
form the Hudsonian forest, discontinuous because of the tremendous 
rock slides and glacial cirques. Tuolumne Meadows, at 8,600 feet, is 
pure Hudsonian and here are found Behr’s Sulphur {Colias hehrii) , 
the Podarce Blue (Plebejus glandon podarce), and the Mariposa Cop- 
per {Lycaena mariposa) . The upper limits of Hudsonian Zone are 
defined by timberline at approximately 11,000 feet. 

Arctic- Alpine Zone: The Alpine Willow, stunted to a height of a 
few inches, and the Sierra Nevada Rosy Finch inhabit the bleak talus 
slides and snow banks above timber line. The zone is characterized by 
perennial herbs with small tops and large root systems. The heathers, 
Bryanthus and Cassiope, and Alpine Sorrel (Rumex) cling to meager 
patches of damp soil. The Ivallda Arctic {Oeneh chryxus ivallda), 
Malcolm’s Checker-Spot {Melitaea damoetas malcolmi) , and the tiny 
Yosemite Blue {Plebejus shasta comstocki) fly from 11,000 feet (occa- 
sionally less) to the tops of Mt. Dana and Mt. Lyell. Recently Miriam’s 
Skipper {Hesperia miriamae) has been added to the strictly Arctic- 
Alpine list. 

It should be remembered that it was in California, and particularly 
on the west slope of the Sierra Nevada, that most of the detailed work 
on life zones, including the delineation of indicator species, was done. 
It is also on the west slope of the Sierra Nevada and Cascade ranges 
that life zones attain their optimum development, occurring in regular 
succession from west to east in response to moderate gradients of 
slope and a comparatively mild, ocean-controlled climate. Elsewhere 
life zones are likely to occur haphazardly, with extreme local conditions 
often overriding the effects of latitude and altitude, which elsewhere 
are the principal factors governing temperature, on which the life- 
zone concept was originally based. One of these, the humidity factor, 
was taken into consideration by Grinnell, Storer, and Dixon in both 
the Yosemite and Lassen sectors, which included an arid region east 
of the Sierra divide. Their concept of a trans-montane fauna con- 
ditioned by high aridity as opposed to a cis-montane fauna con- 
ditioned by high humidity leads us to the principle of replacement of 
one set of zonal indicators by another whenever certain natural boun- 
daries are transgressed. This is better expressed within the context of 
a more recent ecologic concept, that of the Biotic Province. 


2(i):t-^6, 1963 


YOSEMITE BUTTERFLIES 


13 


BIOTIC PROVINCES 

The replacement of one set of indicator species by another can be 
so complete that we may question whether we are dealing with the 
same life zone in two widely separated regions, or even in adjacent 
regions, when each indicator species has been replaced by a different 
but closely related species or subspecies. The solution to this predica- 
ment is found in the concept of the Biotic Province, originated by 
Vestal (1914) and developed by Dice (1943), who divided the North 
American continent into 28 biotic provinces, of which 14 occur in 
states west of the Rockies, 7 in California. These are geographical 
areas, permanent and mutually exclusive, and containing within their 
confines vertical bands that Dice calls, not life zones^ but life belts. In 
such a system the principle of replacement need not be considered, for 
by definition no life belt extends beyond its province, although similar 
life belts may occur in adjacent provinces. 

In the Yosemite region, as has been mentioned, all life zones are 
represented from Lower Sonoran (in the San Joaquin Valley) to 
Arctic-Alpine, with a correspondingly wide spectrum of plant and 
animal life. Park boundaries are drawn, however, in such a manner that 
the eastern boundary coincides with the crest of the Sierra Nevada, 
thus limiting the park proper to one biotic province, the Californian. 
If we extend the Yosemite sector eastward to include the eastern slope 
of the Sierra (as did Grinnell and Storer in extending their Yosemite 
sector to the shore of Mono Lake), we will have added another biotic 
province, the Artemesian, and with it the replacement species of that 
province, potentially doubling the list of species for zones above 
6,500 feet. No more than this could be accomplished by combining the 
species lists from two different parks, the one completely Californian, 
the other Artemesian. Furthermore, such an extension is necessary in 
order to compare the butterfly faunas of Yosemite and Lassen, whose 
boundaries stop at the Sierran Crest, with those of Glacier and Rocky 
Mountain, whose boundaries include territory on both sides of the 
Continental Divide. 

An unexpected dividend for the biologist of this meeting of faunas 
along the border of two biotic provinces is the opportunity of studying 
natural hybridization. Perhaps no better experimental material has 
been provided by nature for this purpose than the butterflies known 
as Admirals (genus Limenitis') . These are found throughout North 
America and are known as allopatric, or replacement, species. Oppor- 
tunity to observe natural hybridization between Californian L. lorquini 
and Artemesian L. weidemeyerii nevadae occurs where their ranges 
meet on the west shore of Mono Lake. A similar opportunity to observe 
natural hybridization between Montanian L. lorquini hurriaoni and 
Saskatchewan L. arthemis rubrofasciata occurs on the east side of 
Glacier Park, at Many Glacier. Whether the fact that they hybridize 
readily reduces the Limenitis complex to a single polymorphic species 


14 


GARTH AND TILDEN 


/. Res. Lepid. 


is a debatable question; however, it should be noted that the interbreed- 
ing is limited to a narrow buffer zone and does not extend to entire 
populations, and that wherever it occurs both parent species are pres- 
ent, as would be required to perpetuate the hybrid, which is presumed 
to be sterile. 

PLANT COMMUNITIES 

A more subtle ecologic division than either life zone or biotic prov- 
ince has long been sought by entomologists attempting to define more 
closely the habitats of insect species. This has at last been provided 
in the system of plant communities developed by Munz and Keck 
( 1949 , 1950 ). Of the 28 California plant communities recognized by 
these authors, at least 16 (plus two of our own designation) occur in 
the greater Yosemite region. Here an attempt is made to list in se- 
quence the plant communities that one might encounter in crossing the 
Sierra Nevada from San Joaquin Valley to Mono Lake, with a side trip 
into Yosemite Valley: 

1. Valley Grasslands (on leaving the Valley) . 

2. Foothill Woodland ( as around Cathay) . 

3 . Chaparral (the familiar "elfin forest”). 

4. Coastal Sage Scrub (formerly California Sagebrush Associa- 
tion, includes Honey Sage Scrub, occurs sporadically). 

5. Oak Woodland (including Golden Cup Oak lands; as Yose- 
mite Valley). 

6. Mixed Evergreen Forest (Laurel-Madrono association; as 
Yosemite Valley walls). 

7. Yellow Pine Forest (monotypic, with an understory of 
"Mountain Misery”). 

8. Mixed Forest (non-pure Douglas Fir, Yellow Pine, and broad- 
leaf trees below pure conifers). Our designation; probably 
an ecotone. 

9 . White Fir-Cedar Forest (on cut-over land, as at Mather). 
Our designation; a successional sub-climax or post-climax 
community. 

10. Red Fir Forest (as between Tamarack Flat and Smoky Jack). 

11. Lodgepole Pine Forest (as Tuolumne Meadows). 

12. Subalpine Forest (often with Mountain Hemlock and White- 
Bark Pine; as Tioga Pass) . 

13 . Alpine Fell-Fields ("rock gardens” above timber line; as 
Gay lor Lakes). 

14. Juniper Woodland (eastern declivity; as below Tioga Pass). 

15 . Pihon- Juniper Woodland (lower down; as around Lee Vin- 
ing). 

16. Sagebrush Scrub (Great Basin Sage, Artemesia tridentata) . 

17. Shadscale Scrub (the Salt-Bush type of scrub). 

18. Alkali Sink (as around Mono Lake) . 


2(t):i-^6, 1963 


YOSEMITE BUTTERFLIES 


15 



Figure 2. Map showing the YOSEMITE high country, indicating roads . trails. .......... 

streams , areas over 11, 000 feet in elevation and collecting localities: 

1933-1934 : 3, Mt. Dana; 6, Florence Lake; 8, Glen Aulin; 9, Kuna Crest; 12, Mt Lyell; 18, Cold- 
water Canyon. 

1956-1962; 59 , Agnew Pass; 60, Bert L.; 61, Cockscomb Pk. ; 62, Mt. Dana, N. slope; 63, Dog L. ; 

64, Donohue Pass; 65, Elizabeth L.; 66, Gaylor L. ; 67, Glen Aulin; 68, Helen L. ; 

70, Lembert Dome; 71, Lyell Base Camp; 72, Lyell Fork Meadow; 74, Mammoth Pk. ; 
75, Merced R.; 76, Mono Pass; 77, Pilot Pk. ; 78, Rafferty Ck. Tr.; 79, Saddlebag 
80, Tioga Pass; 81, NW above Tioga Pass; 83, Thousand Island L.; 84, Tuolumne 
Meadow; 85, Unicorn Pk.; 86, Vogelsang Pass. 

East slope , 1956-1962: 92, Gull L.; 93, June L.; 94, LeeVining Cr. ; 95, Mono L. ; 98, Warren Ck. 


16 


GARTH AND TILDEN 


/, Res. Lepsd. 


[One of US (Tilden) suggests the presence of still another 
east-slope brush community, consisting of Cercocarpus, Pur- 
shia, and desert Pr7mus, not provided for in the system pro- 
posed by Munz and Keck.] 

It will be noted that by far the greater number of plant communi- 
ties ( 1 3 as against 5 ) are found on the western slope, which has the 
greatest extent both in distance and in elevation. It will further be 
seen that as one progresses beyond 13, which represents the Sierran 
crest, the plant communities are entirely different from those before 
13, yet they occupy corresponding altitudinal belts and life zones in 
reverse order. This difference in plant life is paralleled by a difference 
in animal life, including the insects, and is evidence that each slope lies 
in a different biotic province, where relative humidity, rather than 
temperature difference based on altitude, is the controlling factor. Per- 
haps no better example could be given of the need for both systems 
(life zones and biotic provinces) . 

FOOD PLANTS 

The lepidopterist who pursues his interest seriously becomes in- 
creasingly aware of the dependence of insects on plants. For not only 
do butterflies as adults seek flowers for nectar, but as caterpillars they 
depend on vegetation for sustenance. Many are highly selective in their 
choice of plant food, adhering to one kind and rejecting all others. 
More frequently, however, any one of a group of related plants will 
serve: witness the predilection of the Whites for mustards, or of the 
Sulphurs, for legumes. The lepidopterist soon learns to equate these 
butterfly preferences with plant groups possessing certain structural 
resemblances, particularly as regards their flowers, and recognized by 
botanists as plant families. A list of thirty-eight common plant families 
and one hundred eleven of their genera on which butterflies of 
the Yosemite region are known or believed to feed as larvae is pre- 
sented with the hope that it will lead to further investigation. 

Occasionally, and for no apparent reason, a butterfly will be found 
to feed upon a plant quite unrelated to those usually preferred by its 
group. Thus Colias hehrii is known to be a Vaccinium feeder, whereas 
most .species prefer legumes. But as Hovanitz (1950) has 

shown, the V accinmm-iGQ6.mg habit is shared by several Colias species 
in other parts of the world. Since it is highly improbable that such a 
choice of food plant would have been made independently by each of 
them, this food habit becomes a clear indication of relationship, setting 
the Vaccinium feeders apart from others of their genus. When one of the 
Whites, Neophasia menapia, forsakes the mustards usual to its group 
for the needles of a pine tree, we may be certain that it is not tempor- 
arily off its diet, but that it is following a behavior pattern as old as the 
genus itself, and perhaps responsible for its segregation. 


2(i):i-9S, 196} 


YOSEMITE BUTTERFLIES 


17 


Often a butterfly named for the plant believed to be its host has 
proven to feed upon another. The hair-streak, Strymon adenostomatis, 
was found to feed on Mountain Mahogany (Cercocarpus) , and not on 
Chamise {Adenostoma) , as its given name would imply. (It should 
have been called cercocarpivofous”\ ) Indeed, the question as to wheth- 
er the Blues, Plebejus acmon 3.nd P. lupini, deserve specific distinction 
may ultimately rest upon whether they feed exclusively on Lupine 
(Lupinus) and on Buckwheat {Eriogonum) , respectively, or indis- 
criminately on both. 

Yosemite butterflies for which the larval food plant is either un- 
known or in need of confirmation are the following: 


Unknown 

Melitaea damoetas malcolmi 
Melitaea hoffmanni 
Callophrys doudoroffi windi 
C allop hr ys lemberti 
Lycaena nivalis 
Plebejus glandon podarce 
Plebejus shasta comstocki 
Lycaeides ar gyro gnomon anna 
Thorybes mexicana nevada 
Thorybes diver sus 
Erynnis persius 
Erynnis propertms 
Polites sonora 


Needing Confirmation 

Callophrys nelsoni (Libocedrus?) 

Lycaena rubidus ( Rumex ? ) 

Lycaena phlaeas hypophlaeas 
( Rumex ? ) 

Plebejus lupini (Eriogonum?) 

Philotes speciosa {Eriogonum 
in Yosemite.^) 

Colias zerene {Amorpha not 
found in Yosemite) 

Parnassius clodius baldur 
(for Vaccinium only) 

Pyrgus ruralis ( for Potentilla 
— Horkelia only) 

Erynnis pacuvius lilius 
( Ceanothus ? ) 


A documented observation of oviposition (egg laying) of any of 
these species, with positive identification of the plant of choice, or 
better still, the rearing of any of them from egg to adult would be an 
important contribution to the natural history of the Yosemite region. 


SOME PLANTS ON WHICH YOSEMITE BUTTERFLIES 
FEED AS LARVAE 

In the left hand column of the list ( Table 1 ) that follows, host 
plants are listed alphabetically by family and genus. (To have listed 
plants by species would have prolonged the list unduly.) In the 
middle column the common name of the plant is given. In the right 
hand column butterfly species known or believed to feed on that 
plant are listed alphabetically by genus and species. (Subgenera are 
omitted, as are subspecies, where more than one occurs in Yosemite, 
unless each is known to have a different food plant.) The symbols o, 1, 


18 


GARTH AND TILDEN 


/. Res. Lepid. 


and r indicate that the butterfly has been seen to ovipost on, the larvae 
have been taken from, or the insect reared to maturity on that plant 
by one of us. Otherwise, names of host plants have been taken from 
the literature, carefully culled to eliminate genera not known to occur 
in the Sierra Nevada. Butterfly species feeding on more than one plant 
are starred*. 

Grasses (POACEAE=GRAMINACEAE) and Sedges (CYPER- 
ACEAE) are not listed because detailed information concerning the 
genera involved is lacking. Known to be grass feeders are all the 
SATYRIDAE and, of the HESPERIIDAE, the subfamily Hesperiinae. 
It is believed that when this information becomes available, it will be 
shown that butterflies exhibit as marked a preference for certain of 
the monocotyledonous grasses and sedges as they do for certain of the 
dicotyledonous plants. 

THE GEOLOGICAL BACKGROUND 

The landscape of the Yosemite region has been moulded by many 
forces, but by far the most impressive of these has been glaciation. 
The work of glaciers may be observed in the U-shaped gorges of the 
Tuolumne and Merced rivers, in the hanging valleys of Little Yosemite 
and Bridal Veil Creek, in the polished domes and erratic boulders, in 
the glacial lakes (called tarns) and cirques, and in the minaret summits 
of Cathedral and Unicorn peaks. Less evident, but no less genuine, are 
the effects of glaciation on the flora and fauna, including the butter- 
flies. For as the glaciers retreated, the cold-adapted species followed 
them northward and to the mountain tops, where small glaciers con- 
tinue to exist on the shaded slopes of Lyell, Maclure, Dana, and Con- 
ness. Thus the entire upper tier of life zones, the Canadian, Hudsonian, 
and Arctic-Alpine, is inhabited by butterfly species whose nearest rela- 
tives fly either on other isolated mountain tops in temperature regions 
or at near sea level in Canada, Hudson Bay, or the Arctic regions, as 
these zonal names imply. This discontinuous "boreo-alpine” distribu- 
tion is as much a consequence of Pleistocene glaciation as the polished 
granite over which one walks along the Merced Lake trail, or the 
lateral moraines that rim the montane meadows. 

A second consequence of Yosemite’s geological history is that those 
butterflies that now fly above timberline, and in particular the grass- 
feeding Satyridae, such as The Ivallda Arctic {Oeneis chryxus ivallda) 
and Riding’s Satyr {Eumenis ridingsii) , and Hesperiinae, such as the 
newly found Miriam’s Skipper {Hesperia miriamae) , having no trees 
but only barren rock on which to alight, have come to resemble exactly 
the color, shade, and texture of the particular rock found in their re- 
stricted habitat, whether it be sedimentary or metamorphic. (See 
Hovanitz, 1940). In order that the lepidopterist may better appreciate 
these earthen pigments that provide the backdrop for the Yosemite 


1963 


YOSEMITE BUTTERFLIES 


19 


landscape and constitute the palette from which the colors of Yose- 
mite’s alpine butterflies (Plate III) have been selected, although with- 
out design on the insect’s part, the following quotation by Francois E. 
Matthes, father of Sierran geology, is given. The italics are the authors’. 

"Next to the broad belt on the lower slope of the Sierra Nevada, the 
masses of metamorphic rocks situated near the crest of the range are the most 
extensive. They make up the bulk of Mount Dana, Mount Gibbs, and Parker 
Peak, as well as the jumble of mountains north of Tioga Pass, whose central 
summit is Mount Warren. These mountains, in consequence of their compo- 
sition, are variously tinted in subdued yellotvs, browns, reds, and purples and 
by contrast with the pale-gray peaks of granite near by appear somber, as if 
overcast by perpetual shadows. The metamorphic rocks in this crest region, 
however, differ appreciably in character as well as in structure from those 
in the lower belt. There is but little slate among them, schist, quartzite, and 
volcanic rocks being predominant; and the folding is less deep and less 
complex. These facts are readily observed on and about Mount Dana, which 
is capped by gently flexed beds of volcanic origin.” 

— Matthes, 1930, p. 26. 


ACKNOWLEDGMENTS 

Among agencies and individuals cooperating in the preparation 
of this report should be mentioned the National Park Service, Depart- 
ment of Interior, and in particular Park Superintendent John C. Pres- 
ton and Chief Park Naturalist Douglass H. Hubbard, Yosemite 
National Park, who issued the necessary permits, and Park Naturalist 
Keith Trexler, who assisted with the collecting. Among those provid- 
ing suggestions and information on technical matters were John Burns, 
Harry Clench, William Hovanitz, C. Don MacNeill, Paddy McHenry, 
and Paul Opler. Among those furnishing advance information concern- 
ing manuscripts in preparation were Cyril F. dos Passos, C. Don 
MacNeill, and Paul and Anne Ehrlich. Among museums and curators 
whose collections were consulted were the California Academy of 
Sciences, Edward S. Ross, and the Los Angeles County Museum, Lloyd 
M. Martin. Finally, two investigators whose efforts so splendidly suj>- 
plemented our own, and whose records and observations we have used 
to such advantage, were Edmund D. Godwin and Allan Oakley Shields. 

REGARDING NAMES USED IN THIS PAPER 

Since the publication of "Butterflies of California” (Comstock, 
1927), no comparable book on California butterflies has appeared. 
The excellent color plates make this a most useful reference, but time 
and the activities of butterfly students have brought about many 
changes. Additional species have been found to occur in California; 
numerous subspecies have been described; and many revisionary works 
have appeared. Among these may be mentioned those on Ringlets 
(Davenport, 1941; Brown, 1955), on Fritillaries (dos Passos and 


20 


GARTH AND TILDEN 


/. Res. Lepid. 



Fig, 3. Mt. Lyell, el. 13,900 ft., from tree-line at about 11,000 ft. 
(Upper Lyell Base Camp). Arctic-Alpine life zone, Alpine Fell-Fields, with 
montane meadow and rock garden associations. Habitat of Lycaena phlaeas 
hypophlaeas, Euphydryas editha nubigena, Melitaea damoetas malcolmi, and 
Oeneis chryxus ivallda. — Joseph S. Dixon. 


2( i ); i -96, 1963 


YOSEMITE BUTTERFLIES 


21 


Grey, 1947), on Checker-Spots (Gunder, 1929; Bauer, 1961), on 
Metal-Marks (Opler and Powell, 1961), on Hair-Streaks (Ziegler, 
I960; Clench, 1961), on Coppers (Klots, 1936; Clench, 1961), on 
Blues (Nabokov, 1949; Downey, 1961), and on Skippers (Lindsey, 
Bell, and Williams, 1931; Lindsey, 1942). As a result, only a fraction 
of our California butterflies continue' to bear the generic and specific 
names by which they were known as recently as 1927. 

Following the publication of "Check List of Lepidoptera of Canada 
and the United States of America” (McDunnough, 1938), incorpor- 
ating changes recommended in "Generic Names of Holarctic Butter- 
flies” (Hemming, 1934), together with the findings of many other 
workers, no similar standard of reference has been advanced. Publica- 
tion of a new check list by C. F. dos Passos is believed imminent, and 
changes made since 1938 will be reflected therein. Many of these will 
have been anticipated in "A Field Guide to the Butterflies” (Klots, 
1951 ), but this admirable little book treats only the eastern species. 
Still other modifications appear in "How to Know the Butterflies” 
(Ehrlich and Ehrlich, 1961 ) . 

In deciding what names to use in this article, the authors have tried 
to steer a moderate course, adopting many names now accepted by 
common usage, but avoiding some of the more recent innovations until 
these shall have met with general approval. The sequence and arrange- 
ment of genera and families follows as closely as possible what we are 
led to believe will be the arrangement of the forthcoming check list 
already referred to. The more advanced student as well as the pro- 
fessional should be able to make use of that publication in conjunc- 
tion with this one, which is intended primarily as a field guide, and 
not primarily as a contribution to the systematic literature. 

The authors’ departure from a strict interpretation of the Rules of 
Zoological Nomenclature in designating the western subspecies of 
AtUdes halesus as estesi Clench rather than as corcorani Gunder is de- 
liberate. For while it is conceded that Gunder’s name is the older of 
the two, it was applied, not to the entire race, but to a single aberrant 
individual. We believe that intent should be given due weight in such 
matters, and it was certainly not Gunder’s intent to define the western 
subspecies, whereas it was Clench’s. The same logic is followed in 
attributing Lycaeides melissa inyoensis to Nabokov, and not to Gunder. 


22 


GARTH AND TILDEN 


/. Res. Lepid. 


ACCOUNT OF SPECIES OF THE YOSEMITE REGION 

SATYRIDAE 

la. Coenonympha tullia californica West. & Hew. PI. I, fig. c 
b. Coenonympha tullia mono Burdick PI. IV, fig. a 

THE CALIFORNIA RINGLET and THE MONO RINGLET 
are the two races of this widespread species found in the Yosemite 
region, the former flying in the western foothills, the latter on the 
sage-brush flats of Mono Basin. Weak fliers, they keep close to the 
grasses on which the larvae feed. Following the studies of Davenport 
(1941) and Brown (1954, 1955), all North American ringlets except 
the rare Hayden’s Ringlet, Coenonympha haydenii (Edw.), of Wy- 
oming and adjacent states are considered subspecies of the Eurasian 
C. tullia (Muller), although Burdick (personal communication) was 
inclined to regard his mono as a derivative of the Great Basin-Rocky 
Mountain C. t. ochracea Edw., which he appeared to regard as a dis- 
tinct species. 

Life Zones: Upper Sonoran (Transition). Plant Communities: Val- 
ley Grasslands, Foothill Woodland; Sagebrush Scrub, Marsh. Host Plants: 
POACEAE (Grasses). 

2. Cercyonis pegala ariane (Bdv.) 

It is with some hestitation that the name THE ARIANE SATYR 
is applied to specimens from Mono Lake, because it is BARON’S 
SATYR, Ceifcyonis pegala haroni (Edw.), that occurs in a similar 
situation in northeastern California and southeastern Oregon. 

Life Zones: Upper Sonoran. Plant Communities: Pihon-Juniper 
Woodland, Marsh, and Montane Meadow. Host Plants: POACEAE 
(Grasses). 

3. Cercyonis silvestris (Edw.) PI. I, fig. e 

THE SYLVAN SATYR is a creature of the chaparral forest that 

clothes the foothills. A lover of shade, it seeks to lure the pursuer ever 
deeper into the undergrowth. Its favorite haunts are groves of Man- 
zanita. Scrub Oak, and Mountain Mahogany, interspersed with an occas- 
ional Digger Pine. 

Life Zones: Upper Sonoran (Transition). Plant Communities: 
Foothill Woodland, Chaparral. Host Plants: POACEAE (Grasses). 

4. Cercyonis oeta (Bdv.) PI. IV, fig. c 

THE LEAST SATYR was not encountered in Yosemite Park 

during the 1933-1934 period, although it was known to inhabit the 


i$6j 


YOSEMITE BUTTERFLIES 


23 


dry eastern slope of the Sierra Nevada. Since that time, however, it 
has been found on several occasions in the Tioga Pass region, but 
always within sight of the Sierran Divide. 

Life Zones: Transition, Canadian, Hudsonian ( Arctic- Alpine ) . 
Plant Communities: Juniper Woodland, Pihon-Juniper Woodland, 
Sagebrush Scrub (Alpine Fell-Fields). Host Plants: POACEAE 
(Grasses) . 

5. Oeneis chryxus ivallda (Mead) PI. Ill, fig. b 

THE IVALLDA ARCTIC inhabits the bleak, wind-swept sum- 
mits that culminate in Mt. Lyell, Mt. Dana, and Mt. Conness. They 
may be encountered in a boulder-strewn clearing between stands of 
Mountain Hemlock, as near Vogelsang Camp, on barren ridges, as the 
crest above Tioga Pass crossed by the Gaylor Lakes trail, or on the 
moraines just east of Tioga Pass. Edmund Godwin reported "many 
seen at 12,000 feet on Matterhorn” in 1934. Erratic fliers, extremely 
difficult to capture, the Arctics have a habit of leaning with the wind 
when alighting, their mottled wings resembling a patch of lichen. 
According to Hovanitz (1940), the gray forms are found on granite, 
the brown forms on sedimentary rock. The line of contact between 
the two formations passes through the Yosemite sector and is a con- 
spicuous feature of Sierran geology. Because of the inaccessibility of 
its habitat, the IVALLDA ARCTIC presents the ultimate challange to 
the energetic lepidopterist. 

Life Zones: Arctic- Alpine (Hudsonian). Plant Communities: Al- 
pine Fell-Fields, Subalpine Forest (Glacial Moraine). Host Plants: 
POACEAE ( Grasses ) . 

DANAIDAE 

6. Danaus (Danam) plexippus (Linn.) 

THE MONARCH is common in Yosemite Valley and throughout 
the lower elevations of the park. Various species of Milkweed grow 
along the Merced River at Old Village, at Mirror Lake, and at the 
fork in the road to the Giant Yellow Pine, as well as along the higher 
trails that follow the canyon rim. It also occurs on the eastern Sierra 
slope, at Mono Lake, and has been found at Upper Lyell Base Camp 
( Shields ) . The banded larvae are easily discernible, but the waxy green 
chrysalids defy detection. 

Life Zones: Unrestricted below Hudsonian. Host Plants: ASCLE- 
PIADACEAE: Asclepias (True Milkweeds). 

7. Danaus (Tasitia) gilippus strigosus (Bates) 

THE STRIATED QUEEN in another milkweed feeder that occurs 
in Yosemite territory only on the dry east side of the Sierra Nevada, 
in the vicinity of Mono Lake. It may be distinguished from Danaus 
{Tasitia) gilippus berenice (Cram.) of the southeastern United States 
by the scattered white scales along the wing veins. Both are sub- 
species of the tropical D. (T. ) gilippus (Cram.), ranging northward 
from Brazil. 


24 


GARTH AND TILDEN 


/. Res. Lepid. 


Life Zones: Sonoran, both Lower and Upper (= Austral). Plant 
Communities: Many. Host Plants: ASCLEPIADACEAE: Asclepias 
(True Milkweed), and several other milkweed genera. 

NYMPHALIDAE 

S'. Argy finis {Semnopsyche) cybele leto ISeht: 

THE LETO ERITILLARY is the handsomest butterfly of the 
Yosemite Valley and the species most certain to attract the attention 
of the park visitor in the late summer season. Erom the thistles at 
Mirror Lake, which they visit for nectar, the butterflies may be traced 
to the sequestered meadow near Camp 9. Here the velvet-brown females 
deposit their eggs among the dried grasses, while the ruddy males hover 
above them, for leto is one of the two California fritillaries in which 
the sexes differ in color, the other being the following apacheana. 

Life Zones: Transition. Plant Communities: Poplar Woodland 
(Black Cottonwood). Host Plants: VIOLACEAE: Viola (Violet). 

9. Argynnis {Speyeria) nokomis apacheana Skin. 

PL IV, fig. k 

THE APACHE ERITILLARY, athough of another species, may 
be considered the eastern Sierran counterpart of The Leto Eritillary of 
the western Sierra. Even more spectacular than leto, it flies in the well 
watered meadows of the Owens River drainage, and has been found 
recently at Gull Lake and Mono Lake. The females emerge a full two 
weeks later than the males, and so dissimilar are the sexes in color 
that they might be mistaken for different species. The under side of 
the male is illustrated in color. 

Life Zones: Upper Sonoran (Transition). Plant Communities: Wet 
Meadow, Marsh. Host Plants: VIOLACEAE: Viola (Violet). 

10a. Argynnis {Speyeria) zerene zerene Bdv. PL I, fig. b 

b. Argynnis {Speyeria) zerene malcolmi Comst. 

THE ZERENE ERITILLARY is common along Highway 41 from 
Chinquapin to Wawona, in the Mariposa and Tuolumne Big Tree 
groves, and at Mather and Hetch-Hetchy summit. Every patch of 
thistle will bear investigation, for this fritillary is highly susceptible 
to its lure. Its Sierran east-slope counterpart, MALCOLM’S FRIT- 
ILLARY, is found along the road from Highway 395 to Mammoth, 
and at Warren Creek on the Tioga Road. Its first choice for nectar is 
Monardella, a dwarf mint. 

Life Zones: Transition, Canadian. Plant Communities: Yellow 
Pine, White Fir (Red Fir), Mixed Coniferous forests for A. z. zerene, 
Sagebrush Scrub, Jeffrey Pine Forest, Pihon-Juniper, and Juniper 
Woodland for A. z. malcolmi. Host Plants: VIOLACEAE: Viola 
(Violet). 

11a. Argynnis {Speyeria) callippe inornata Edw. 

b. Argynnis {Speyeria) callippe nevadensis Edw. PL IV, fig. b 
THE PLAIN ERITILLARY flies in the western foothills of the 
Yosemite region in the late spring. It has been found at Fish Camp, 


2(t):i‘p6, 196} 


YOSEMITE BUTTERFLIES 


25 



Fig. 4. Research Reserve, el. 8,700 ft. Open slope with Western White 
Pine predominant; Lodgepole Pine and Red Fir also present, with understory 
of Manzanita and Huckleberry Oak. Canadian life zone. Philotes battoides 
and P. shasta comstockii hover over the dwarf buckwheat and sparse grasses 
ruralis may be seen here.—Neva Snell. 



26 


GARTH AND TILDEN 


/. Res. Lepid. 


at Briceburg, and at Indian Flat, below El Portal. THE NEVADA 
FRITILLARY, its eastern Sierran counterpart, flies in the Mono Lake 
region in early summer. It has been found at the summit above Crowley 
Lake on Balsam Root ( Balsamorhiza ) , a large-leaved composite. The 
Great Basin races of callippe tend to be green and silvered below, the 
races of the western foothills brown and unsilvered. 

Life Zones: Upper Sonoran (lower Transition). Plant Commun- 
ities: (Chaparral) Foothill Woodland, Yellow Pine and Mixed Forest 
for A. c. inornata; Sagebrush, Pihon-Juniper Woodland, Jeffrey Pine 
Forest openings. Juniper Woodland for A. c. nevadensis. Host Plants: 
VIOLACEAE: Viola (Violet). 

12. Argy finis {Speyeria) egleis Behr 

Syn. Argynnis {Speyeria) montivaga Behr 

THE EGLEIS FRITILLARY, known for many years as THE 
MOUNTAIN VAGABOND, is a species of uniformly small size 
characterized by a slight thickening of the veins of the primaries. It 
flies with The Arge Fritillary at Tioga Pass and is common at all ele- 
vations above 6,000 feet, preferring the minted slopes, whereas A. 
mormonia arge prefers the meadows. On the west slope of Mt. Dana 
it fairly swarms in the late afternoon sunlight. 

Life Zones: upper Transition, Canadian, Hudsonian (Arctic-Al- 
pine). Plant Communities: Openings in Mixed Coniferous, White 
and Red Fir, and Lodgepole Pine Forests (Alpine Fell-FieldsJ . Host 
Plants: VIOLACEAE: Viola (Violet). 

13. Argynnis {Speyeria) atlantis irene Bdv. PL 11, fig. h 

THE IRENE FRITILLARY of Yosemite, as determined for the 

earlier survey by Dr. J. McDunnough, is a dark race, some specimens 
of which are strongly suggestive of The Hydaspe Fritillary. Irene, 
however, is found at elevations well above those at which hyaspe 
occurs, and for this reason should not be confused with it. One of us 
(Tilden) has taken it also on the Sonora Pass Road, between Straw- 
berry and the Clark Fork Road. In Yosemite it is something of a rarity. 

Life Zones: upper Transition, Canadian. Plant Communities: Yel- 
low Pine-Sugar Pine, White Fir and Red Fir Forests. Host Plants: 
VIOLACEAE: Viola (Violet). 

14. Argynnis {Speyeria) hydaspe Bdv. 

THE HYDASPE FRITILLARY first appears in Yosemite Valley 
about the Fourth of July, flying rapidly across the valley floor but paus- 
ing over the Monardella of the talus slopes. It is completely unsilvered 
and the creamy spots on the under side give a "checker-board” effect 
quite different from the purplish shades and frequent light silvering of 
the somewhat similar but usually larger zerene. 

Life Zones: Transition, lower Canadian. Plant Communities: Yel- 
low Pine, Mixed Coniferous, and White Fir Forests. Host Plants: 
VIOLACEAE: Viola (Violet). 


I $6} 


YOSEMITE BUTTERFLIES 


27 


15. Argynms (Speyeria) mormonia arge Stkr. PI. Ill, fig. 1 
The Sierran race of THE MORMON FRITILLARY flies with THE 

EGLEIS FRITILLARY throughout the park. Males lack the enlarged 
veins of the forewing; females are more finely marked than those of 
egleis; otherwise the two species are indistinguishable, and particularly 
so when on the wing. The active males range widely, but the females 
will be found near moist meadows. The Rocky Mountain race of A. 
(S.) mormonia is eurynome Edw. 

Life Zones: Canadian, Hudsonian ( Arctic-Alpine) . Plant Com- 
munities: Sub-alpine Meadows; Alpine Fell-Fields, Meadow and Grass- 
land. Host Plants: VIOLACEAE: Viloa (Violet). 

16. Boloria {Clossiana) epithore (Edw.) PL II, fig. d 

THE WESTERN MEADOW FRITILLARY favors small, open 

glades beneath fir trees. Along Yosemite Creek it is particularly abund- 
ant in mid-July. There is also a colony on Dana Trail, where it passes 
through a marsh before rising sharply onto the slope of the mountain. 
The under side of the secondaries resembles Boloria ( C. ) toddi ( Holl. ) , 
a widely ranging eastern species; however, in epithore the outer margin 
of the primaries is rounded, whereas in toddi it appears square-cut. 

Life Zones: Canadian (Hudsonian). Plant Communities: White 
and Red Fir Forest; Bogs and Marshes. Host Plants: VIOLACEAE: 
Viola (Violet). 

17a. Euphydryas chalcedona chalcedona (Dbldy. & Hew.) 
b. Euphydryas chalcedona sierra (Wgt.) 

THE CHALCEDON CHECKER-SPOT ranges throughout the 
foothills of the western portion of the state, being replaced in the 
Sierran highlands by THE SIERRA CHECKER-SPOT, now recog- 
nized, through the work of J. D. Gunder (1929), as a subspecies, 
or geographical race. Typical chalcedona may be found abundantly 
among the chaparral-clothed foothills, whereas race sierra prefers the 
wooded glades of the evergreen belt. A variety of food plants is 
acceptable to the voracious larvae. 

Life Zones: Upper Sonoran for E. c. chalcedona; Canadian for E. c. 
sierra. Plant Communities: Foothill Woodland, Chaparral for E. c. 
chalcedona; Fir Forest for E. c. sierra. -Host Plants: SCROPHULAR- 
lACEAE: Scrophularia (Bee Plant); Pentstemon ( Pentstemon ) ; 
Castilleja (Paint Brush); Mimuh/s (Monkey Flower); and Diplacus 
(Sticky Monkey Flower) for E. c. chalcedona; BOR AGIN ACE AE: 
Mertensia ciliata var. stomatechoides (Mertensia), for E. c. sierra, ac- 
cording to Davenport and Dethier (1937) CAPRIFOLIACEAE: 
Symphoricarpos albus (Snow Berry) for E. c. chalcedona. 

18a. Euphydryas editha ruhicunda (Hy. Edw.) PL I, fig. d 

b. Euphydryas editha nuhigena (Behr) PL III, fig. c 

c. Euphydryas editha monoensis Gund. PL IV, fig. f 

THE RUDDY CHECKER-SPOT flies in the western Sierra from 


28 


GARTH AND TILDEN 


/. Res. Lepid, 


Tulare to Sierra counties, its "lowland” counterpart of the eastern 
Sierra being THE MONO CHECKER-SPOT. As suggested by Com- 
stock (1927), and confirmed by Gunder (1929), THE CLOUD- 
BORN CHECKER-SPOT, E. nuhigena, has proven to be a delicately 
marked dwarf form that has adapted itself to alpine areas. All are sub- 
species of the widely ranging Etiphydryas editha Bdv., each with a well 
defined and mutually exclusive range of its own. 

Life Zones: Upper Sonoran, Transition for E. e. rubicunda and E. e. 
monoensis; Hudsonian, Arctic-Alpine for E. e. nuhigena. Plant Com- 
munities: Foothill Woodland for E. e. rubicunda; Pihon-Juniper Wood- 
land for E. e. monoensis; Alpine Fell-Fields for E. e. nuhigena. Host 
Plants: PLANTAGINACEAE: Plantago (Plantain). 

19. Melitaea {Chlosyne) damoetas malcolmi Comst. 

PL III, fig. a 

MALCOLM’S CHECKER-SPOT was described from the then only 
known colony at Red Lake, el. 11,000 ft., above the Mammoth Lakes 
Basin, where it flies on a precipitous talus slide. It is now known to 
fly in the Yosemite region and throughout the central Sierra at cor- 
responding elevations. Perhaps the most intimate knowledge of the 
species is possessed by Oakley Shields, who has combed the high coun- 
try for isolated colonies. The most accessible of these is located above 
Upper Gaylor Lake near an abandoned stone cabin reached by a fish- 
erman’s trail from Tioga Pass. 

Life Zones: upper Hudsonian, Arctic- Alpine. Plant Communities: 
Alpine Fell-Fields. Host Plants: Not known. Possibly COMPOSITAE: 
Aster ( Aster ) . 

20. Melitaea {Chlosyne) acastus Ed'w. 

THE ACASTUS CHECKER-SPOT is a Great Basin flier, found 
sparingly around Mono Lake. The few records from late June and 
early July suggest an earlier flight season. 

Life Zones: Upper Sonoran. Plant Communities: Sagebrush Scrub, 
Meadowland. Host Plants: COMPOSITAE: Aster (Aster). 

21a. Melitaea {Chlosyne) palla palla Bdv. PL II, fig. j 

b. Melitaea {Chlosyne) palla whitneyi Behr 

THE NORTHERN CHECKER-SPOT ranges widely throughout 
the Sierra Nevada and Cascade chains as well as the Rockies. It is re- 
placed at higher elevations in the southern Sierra by WHITNEY’S 
CHECKER-SPOT, a darker and ruddier form. Whether whitneyi 
actually occurs in the Yosemite region is a matter for further investi- 
gation. None were encountered in the course of the present survey. 

Life Zones: Upper Sonoran, Transition, Canadian for M. (C.) p. 
palla; Hudsonian for M. (C.) palla ivhitneyi. Plant Communities: 
Foothill Woodland, Chaparral, Mixed Evergreen Forest. Host Plants: 
COMPOSITAE: Aster (Aster). 


2(i)ti-96, 196) 


YOSEMITE BUTTERFLIESi 


29 


22. Melitaea {Chlosyne) hoffmanni Behr PI. II, fig. 1 

HOFFMANN’S CHECKER-SPOT was the most abundant but- 
terfly on the Research Reserve in 1933. They fairly swarmed over 
Monardelh; the nectar seeming to intoxicate, or at least render them 
oblivious to the observer’s presence. In 1956 but a few were seen on 
a north-facing slope along the Tioga Road opposite the White Wolf 
turnoff. 

Life Zones: Canadian. Plant Communities: Openings in Fir For- 
est. Host Plants: Not known. 

23 . Melitaea {Chlosyne) leanira F. & F. 

THE LEANIRA CHECKER-SPOT invades the western portions 
of the park from the foothills east of the San Joaquin Valley, where 
it properly inhabits the chaparral. It is common at Indian Flat, below 
El Portal. 

Life Zones: Upper Sonoran (Transition). Plant Communities: 
California Sagebrush Scrub (also called Coastal Sage Scrub), Foothill 
Woodland. Host Plants: SCROPHULARIACEAE: Castilleja (Indian 
Paint Brush), Cordylanthus tenuis (Bird’s Beak). 

24a. Phyciodes {Phyciodes) campestris campestris (Behr) 
b. Phyciodes {Phyciodes) campestris montana (Behr) 

THE FIELD CRESCENT is found about moist places at lower 
elevations, as on the floor of Yosemite Valley, or at Mather, just outside 
the park. Higher up it grades imperceptibly into THE MOUNTAIN 
CRESCENT, in which the tawny ground color has all but obliterated 
the submarginal row of spots on the secondaries. Although heretofore 
treated as a distinct species, on the basis of Yosemite specimens the 
present authors prefer to consider montana but an altitudinal race of 
campestris. Specimens from Mono Lake are clearly intermediate. 

Life Zones: Upper Sonoran, Transition for P. (P.) c. campestris; 
Canadian, lower Hudsonian for P. (P.) c. montana. Plant Communi- 
ties: Several. Wet Meadowland, Roadside Associations. Host Plants: 
COMPOSITAE: Aster (Aster). 

25 . Phyciodes {Phyciodes) mylitta (Edw.) PL II, fig. c 

THE MYLITTA CRESCENT frequents the streamside and moist 

meadows, often in association with the following species, Polygonia 
satyrus (Edw.). It can be recognized by the bright yellow-brown 
ground color. California specimens are typical, that is to say, darker, 
whereas Rocky Mountain specimens incline toward the lighter races 
Phyciodes ( P. ) mylitta pallida Edw. and thehais ( G. & E. ) . 

Life Zones: Upper Sonoran, Transition, Canadian. Plaht Communi- 
ties: Several, as is frequent with wide-ranging species. Host Plants: 
COMPOSITAE: Carduus (Italian Thistle), Cirsium (Native Thistle), 
and Silyhum (Milk Thistle) . 

26. Polygonia satyrus { Edw. ) 

THE SATYR is a streamside flier that alights on the trunks of 





1963 


YOSEMITE BUTTERFLIES 


31 


Alder, Willow, and Cottonwood, to blend indistinguishably with the 
mottled bark. 

Life Zones: Upper Sonoran, Transition. Plant Communities: Foot- 
hill Woodland, Mixed Coniferous Forest, and others. Riparian Associa- 
tion. Host Plants: URTICACEAE: Urtica (Nettle). 

27. Polygonia faunus rusticus ( Edw. ) 

THE RUSTIC ANGLE-WING occurs principally north of Yose- 
mite, but may be looked for. wherever Azalea grows. Its inclusion in 
the Yosemite fauna is based on an early record of Dr. E. O. Essig. The 
specimen was seen by the senior author at the Yosemite Museum in 
1933. 

Life Zones: Transition. Plant Communities: Several. Moist Wood- 
land, Riparian associations. Host Plants: ERICACEAE: Rhododendron 
(Azalea); SAXIFRAGACEAE: Ribes (Currant, Gooseberry); BETU- 
LACEAE: Alnus (Alder); SALICACEAE: Salix (Willow). 

28. Polygonia zephyrus (Edw.) PL II, fig. k 

THE ZEPHYR replaces THE SATYR {Polygonia satyrus) as the 

common Angle- Wing of the Canadian and Hudsonian zones, its pale 
gray underside serving as a distinguishing feature. Yosemite Creek in 
mid-July is a typical situation for the species. Its food plant, Ribes^ is a 
common shrub in the Subalpine Forest. 

Life Zones: Transition (upper), Canadian, Hudsonian. Plant Com- 
munities: Fir Forest, Lodgepole Pine Forest, Subalpine Forest. Host 
Plants: ERICACEAE: Rhododendron (Azalea); SAXIFRAGACEAE: 
Ribes (Currant, Gooseberry). 

29 . Nymphalis calif ornica {^dv.) 

THE CALIFORNIA TORTOISE-SHELL appears in outbreak num- 
bers every few years throughout the Sierra. The June flights observed 
by the senior author in General Grant (now Sequoia-Kings Canyon) 
National Park in 1922, in the San Bernardino Mountains in 1931, and 
in Yosemite in 1933 were aU in a northerly or northwesterly direction, 
roughly paralleling the axis of the Sierra. However, former Park 
Naturalist C. A. Harwell, while on a glacier-measuring survey, reported 
a remrn flight on Oaober 4, 1933, in which myriads of the inseas 
passed through the highest mountain passes, all heading south. The 
duration of the spring flight is but a few days. It begins at dawn and 
lasts until after sundown. At times the greatest density of the insects 
appears to be at tree-top level; again they come so near the ground that 
they may be knocked down with a hat. A few weeks later the Ceanothus 
is matted with webs of the gregarious larvae. Complete defoliation over 
large areas results. The investigation of such an outbreak was the first 
professional assignment of Dr. H. C. Bryant, co-founder of the Yosemite 
Field School. Observations by Edwards and Behr before 1900 in the Mill 
Valley seaion are mentioned by C. B. Williams in his monograph on 
butterfly migration. One of us (Tilden) has observed this phenomenon 
in Yosemite as recently as 1961. The explanation seems to be that the 
CALIFORNIA TORTOISE-SHELL is a swarming species which, like 


32 


GARTH AND TILDEN 


J. Ket. tepid. 


the lemming, has cycles of abundance followed by a drastic reduaion 
in the population, due to causes as yet not fully understood. 

Life Zones: Unrestricted in flight, although the food plant does not 
occur above Canadian. Host Plants: RHAMNACEAE: Ceanothus, 
particularly C. cordulatus (Snow-Bush), and other smooth-leaved species. 

30. Nymphalis milherti furcillata ( Say ) 

MILBERT’S TORTOISE-SHELL has an inward urge to fly upward, 
and has been seen at the very summit of Mt. Lyell, el. 13,090 ft., the 
highest point in Yosemite. Form subpdlida (CklL) flies with normal 
individuals, although not commonly. The subspecific designation furcil- 
lata (Say) is considered applicable to all western specimens south of 
Oregon. 

Life Zones: Transition, Canadian, straying higher. Host Plants: 
URTICACEAE: Urtica (Nettle). 

3 1 . Nymphalis antiopa ( Linn. ) 

THE MOURNING CLOAK is one of the world s most widespread 
butterflies, occurring in Europe, northern Asia, and Japan. In England 
it is called The Camberwell Beauty. 

Life Zones: Unrestricted. Host Plants: SALICACEAE: Salix (Wil- 
low); Populus (Poplar, Cottonwood); ULMACEAE: Ulmus (Elm). 

32. V anessa atalanta (Linn.) 

THE ALDERMAN or THE RED ADMIRAL is found in Europe 
and Japan, as well as in North America. Few butterflies are easier to 
recognize by sight. 

Life Zones: Upper Sonoran, Transition. Host Plants: URTICA- 
CEAE: Urtica (Nettle), Boehmeria (False Nettle), Parietaria (Pelli- 
tory).- 

33. V anessa virgifiiensis {T>ra,) 

THE VIRGINIA LADY is widely distributed throughout North 
America and may be recognized by the large eye-spots on the under 
surface of the hind wings. 

Life Zones: Upper Sonoran, Transition, Canadian. Host Plants: 
COMPOSITAE: Gnaphalium (Cudweed; Everlasting), Anaphalis 
(Everlasting, Antennaria (Everlasting), Artemesia heterophylla 
(Mugwort) . 

34. V anessa cardui {Unn.) 

THE PAINTED LADY or THE THISTLE BUTTERFLY is prob- 
ably the most cosmopolitan of butterflies. Native to the Northern 
Hemisphere, it has been introduced elsewhere. 

Life Zones: Unrestricted. Host Plants: BORAGINACEAE: Am- 
sinckia (Fiddle-Neck), Crypantha (Nievitas); COMPOSITAE: Car- 
duus (Italian Thistle), Cirsium (Native Thistle), Silybum (Milk 
Thistle) . 

35. Vanessa car ye {Hhn,) 

THE WEST COAST LADY is restricted to the western United 


2(i)ti~s6, i$6} 


YOSEMITE BUTTERFLIES 


33 


States^ where it is more common in back yards and vacant lots than in 
the out-of-doors. 

Life Zones: Unrestricted. Host Plants: MALVACEAE: Malm 
(Mallow), Lavatera (Tree Mallow), Sida (Alkali Mallow), sidalcea 
( Checkerbloom, Wild Hollyhock), Sphaeralcea {—Malvastrum) 

( Apricot Mallow, Bush Mallow ) . 

36. Precis orithya evarete (Cram.) 

Syn. Junonia coenia Hbn. 

THE BUCKEYE may be recognized by the brightly colored eye- 
spots on all wings. The aggressive males have been known to select a 
perch and defend a "territory’'. 

Life Zones: Upper Sonoran, Transition, (lower Canadian). Host 
Plants: PLANTAGINACEAE: Plantago (Plantain) ; SCROPHULARI- 
ACEAE: Mimulus (Monkey Elov/er). 

37. Limenitis {Limenitis) weidemeyerii nevadae (B. & B.) 

PI. IV, fig. h 

THE NEVADA ADMIRAL is a Great Basin race of the Rocky 
Mountain Weidemeyer’s Admiral, A (L.) weidemeyerii (Edw.), 

found at Mono Lake on the eastern side of the Sierra Nevada. At Lee 
Vining, where its range overlaps that of the following species, L. (L.) 
lorquini Bdv., the hybrid L. (L.) fridayi (Gund.) occurs. Since the 
blend is an even one, in which the characteristics of neither species pre- 
dominate, we would prefer to indicate it as L. w. nevadae X T. lorquini, 
rather than as a form of the above. 

Life Zones: Upper Sonoran, Transition. Plant Communities: Several. 
Aspen Grove, Willow Thicket, Riparian Associations. Host Plants: 
SALICACEAE: Populus (Aspen, Cottonwood), Salix (Willow). 

38. Limenitis {Limenitis) lorquini Bdv. PI. IL fig. b 

LORQUIN’S ADMIRAL adheres to the riparian association through- 
out three life zones, seldom straying from the willows and poplars of 
the stream bed. On the wing it is sometimes confused with the follow- 
ing species, but the manner of flight is quite different. 

Life Zones: Upper Sonoran, Transition, Canadan. Plant Communi- 
ties: Several. Aspen Grove, Willow Thicket and Riparian Associations. 
Host Plants: SALICACEAE: Salix (Willow), Populus (Aspens and 
Cottonwoods) . ROSACE AE: Prunus ( Choke -cherry ) . 

39. Limenitis (Adelpha) bredowii calif ornica (Bud.) 

PL I, fig. g 

THE CALIFORNIA SISTER flies with the foregoing species, which 
it superficially resembles, but from which it may be told on the wing by 
the habit of alternating a few rapid beats with a glide in which the 
wings are held slightly below the horizontal level. Unlike LORQUIN’S 
ADMIRAL, however, THE CALIFORNIA SISTER forsakes the stream 
margins for the hot canyon walls and the Golden Cup Oak association. 
No other insect is so likely to attract the attention of the park visitor. 


34 


GARTH AND TILDEN 


/. Res. Lepid. 


Life Zones: Upper Sonoran, Transition, Canadian. Plant Communi- 
ties: Oak Woodland, Mixed Evergreen Forest. Host Plants: FAGA- 
CEAE: Quercus (Oak), especially Quercus chrysolepis (Golden Cup 
Oak, Maul Oak ) . 

40a. Apodemia mormo mormo ( F. & F. ) 

b. Apodemia mormo tuolumnensis Opler & Powell 

THE MORMON METAL-MARK is found at higher elevations and 
on the east Sierran slope. THE TUOLUMNE METAL-MARK is known 
only from Pate Valley in the Grand Canyon of the Tuolumne, the type 
locality. It is "a Sierran population characterized by a great extent of 
orange on the upper side together with reduction of white spots and dark 
under side. So far as is known it is restricted to a population along a 
four mile area of the Pate Valley Trail from about 4500’ to 7000’. It is 
likely a segregate from virgulti, its nearest geographic relative, which 
probably occurs in scattered colonies through the lower foothills to the 
west.” (Opler and Powell, 1961) BEHR’S METAL-MARK, Apodemia 
mormo virgulti (Behr) , is the near relative. 

Life Zones: Upper Sonoran (Transition) for A. m. tuolumnensis; 
to Hudsonian for A, m. mormo. Plant Communities: Chaparral for 
A. m. tuolumnensis; Sagebrush Scrub and Pi non- Juniper Woodland for 
A. m. mormo. Host Plants: POLYGONACEAE: Eriogonum (Buck- 
wheat) , perennial species only. 

LYCAENIDAE 

41. Habrodais grunus (Bdv. ) PL I, fig. h 

BOISDUVAL’S HAIR-STREAK is noted for its crepuscular habits. 

A party of the Yosemite Field School, on its way to the Research Reserve, 
discovered it flying at Columbia Point before 5 a.m. in semi-darkness. It 
continues to hover above the live oaks after the last rays of sunlight have 
left the canyon walls. 

Life Zones: Transition (Canadian). Plant Communities: Oak Wood- 
land, Chaparral. Host Plants: FAGACEAE: Quercus chrysolepis (Gol- 
den Cup Oak, Maul Oak); Castanopsis (Chinquapin) suspected. 

42. Atlides halesus estesi Clench 

THE GREAT BLUE HAIR-STREAK displays a metallic blue of 
dazzling luster on the superior surfaces. Rarely encountered north of 
Tehachapi, it has been found several times in Yosemite since the capture 
by a small boy of the specimen recorded by Garth (1935). The worn 
condition of Yosemite specimens suggests flight from lower elevations, 
perhaps outside the park. 

Life Zones : Upper Sonoran ( Transition, Canadian ) . Plant Com- 
munities: Foothill Woodland. Host Plants: LORANTHACEAE : 
Phoradendron ( Mistletoe ) , male flowers only. 

43. Strymon {Strymon) melinus pudica (Hy. Edw.) 

THE COMMON HAIR-STREAK flies in the Museum garden and 
throughout the lower elevations of the park. It is the western race 
pudica, the only race that flies in California. 


2(i):i-96, 1963 


YOSEMITE BUTTERFLIES 


35 


Life Zones: Upper Sonoran (Transition). Plant Communities: Sev- 
eral. Host Plants: BORAGINACEAE: Cynoglossum (Hound’s 
Tongue). LEGUMINOSAE: Astragalus (Loco Weed, Rattle Weed), 
Hosackia {—Lotus) , Lupin us (Lupine). MALVACEAE: Malva (Mal- 
low) . POLYGON ACE AE: Polygonum (Knot Weed). GUTTIFERAE: 
Hypericum (St. John’s Wort). LABIATAE: Hyptis (Bee Sage). 

44. Strymon {Satyrium) fuliginosum semiluna (Klots) 

PI. IV, fig. i 

THE SEMI-LUNATE GOSSAMER WING is an early season flier 
at Mono Lake. Like Melitaea {'Chlosyne) acastus, it has been found in 
late June, but is probably on the wing a month earlier. 

Life Zones: Upper Sonoran. Plant Communities: Sagebrush Scrub. 
Host Plants: LEGUMINOSAE: Lupinus (Lupine). 

45. Strymon {Satyrium) hehrii (Edw.) PL IV, fig. g 

BEHR’S HAIR-STREAK is characteristic of the dry eastern slopes 

of the Sierra Nevada. An isolated colony also exists on the desert slope 
of the San Bernardino Mountains, in southern California.. 

Life Zones: Upper Sonoran. Plant Communities: Sagebrush Scrub. 
Host Plants: LEGUMINOSAE: Lupinus (Lupine), Lotus {=Hos- 
ackia), Astragalus (Loco Weed, Rattle Weed). ROSACEAE: Purshia 
tridentata (Antelope Brush, Bitter Brush), reported (1963). 

46. Strymon {Satyrium) auretorum (Bdv.) PL I, fig. k 

THE GOLDEN HAIR-STREAK may now be reported authori- 
tatively from the Yosemite region on the strength of specimens found 
during the present survey along Hetch-Hetchy Road north of Mather. 
When compared with specimens of The Nut-Brown Hair-Streak, 
Strymon {Satyrium) auretorum spadix (Hy. Edw.), from Crystal Lake 
in the San Gabriel Mountains of southern California, they proved to be 
noticeably different in hue and marking. Specimens were attracted to 
Syringa, Philadelphus Lewisii var. calif ornicus , to the fragrant blooms 
of which the Theclinae are partial. 

Life Zones: Upper Sonoran. Plant Communities: Chaparral. Host 
Plants: FAGACEAE: Quercus (Oak), especially Q. Douglassii (Blue 
Oak) in the Sierra foothills. 

47. Strymon {Satyrium) saepium (Bdv.) 

THE HEDGE-ROW HAIR-STREAK is abundant on the blos- 
soms of the Buckeye, Aes cuius calif ornicus, in the late afternoon, where 
it flies with Strymon { Satyrium ) auretorum and Euphydryas chalcedona. 

Life Zones: Upper Sonoran. Plant Communities: Chaparral (west 
slope); Sagebrush Scrub (east slope). Host Plants: RHAMNACEAE: 
Ceanothus, especially C. cuneatus (Buck Brush). ROSACEAE: Cerco- 
carpus (Mountain Mahogany) . 

48. Strymon {Satyrium) adenostomatis (Hy. Edw.) 

THE GRAY HAIR-STREAK occurs in the western foothills near 
Jersey dale and along the road to Hetch-Hetchy, above Mather. 

Life Zones: Upper Sonoran. Plant Communities: Chaparral. Host 


36 


GARTH AND TILDEN 


/. Res. Lepid. 


Plants: ROSACEAE: Cercocarpus, especially C. betuloides (Hard Tack, 
Mountain Mahogany). Not Adenostoma (Chamise Bush), as its name 
would imply. 

49 . Strymon {Satyrium) sylvinus (Bdv.) 

THE SYLVAN HAIR-STREAK is found about the wiUows of the 
stream bed, rather than in forest glades, as the name might suggest. 
Specimens from Mono Lake show no great tendency toward race 
desertorum (Grin.), which occurs from Kern County north through 
Owens Valley. 

Life Zones: Upper Sonoran (lower Transition). Plant Communi- 
ties: Several. Riparian Association. Host Plants: SALICACEAE: Sdix 
(Willow). 

50. Strymqn {Satyrium) calif ornica (Edw.) 

THE CALIFORNIA HAIR-STREAK is partial to the flowers of 
Pink Pussy-Paws, Spraguea umbellata, from which it may be plucked 
with fingers or with forceps. It is also attracted to Yerba Santa, Erio- 
dictyon calif ornicum, when it visits the nature garden behind the Yose- 
mite Museum. 

Life Zones: Upper Sonoran (lower Transition) on west slope; 
Upper Sonoran and High (arid) Transition on east slope. Plant Com- 
munities: Chaparral, Oak Woodland (west slope); Sagebrush Scrub, 
Juniper Woodland (east slope). Host Plants: RHAMNACEAE: Ceano- 
thus cuneatus (Buck Brush). ROSACEAE: Cercocarpus (Mountain 
Mahogany). Possibly FAG ACE AE: Quercus (Oak). 

51. Strymon {Satyrium) dry ope (Edw.) 

THE DRYOPE HAIR-STREAK was encountered playing about 
sage at Mono Lake. 

Life Zones: Upper Sonoran. Plant Communities: Sagebrush Scrub 
(east slope) ; Chaparral, Foothill Woodland (coastal California). Ripar- 
ian Association. Host Plants: SALICACEAE: Salix (Willow). 

52. Callophrys {Callophrys) dumetorum (Bdv.) 

THE BRAMBLE HAIR-STREAK is the green hair-streak of 
moderate elevations on the western slope of the Sierra Nevada. At high 
elevations, and on the eastern slope, it is replaced by the following 
species. 

Life Zones: Upper Sonoran. Plant Communities: Chaparral. Host 
Plants: POLYGONACEAE: Eriogonum (Buckwheat), especially E. 
nudum. 

53 . Callophrys {Callophrys) lemberti Tilden PL III, fig. k 
LEMBERT’S HAIR-STREAK is the greenish hair-streak of the 

higher elevations of the park. Its relationships appear to be with 
Callophrys (C.) sheridani Edw. of the Rocky Mountains, in which the 
band of white across the under side of the secondaries is more or less 
continuous. Discovered during the course of this survey, it has been 
named in honor of the pioneer resident of Tuolumne Meadows, John 
Batiste Lembertr 


2(i):i-96, 1963 


YOSEMITE BUTTERFLIES 


37 


Life Zones: Hudsonian, Arctic- Alpine. Plant Communities: Alpine 
Fell-Fields. Associated with "rock garden” Eriogonum, E. ovalifolium. 
Host Plants: Unknown. 

54. Callophrys {Mitoura) spinetorum (Hew.) 

THE THICKET HAIR-STREAK is a highly prized find that may 
be recognized by the bold white line across the under side of the second- 
aries. Its dull bluish upper side distinguishes it from the following 
Callophrys {Mitoura) johnsoni (Skin.). 

Life Zones: Transition, Canadian. Plant Communities: Mixed Coni- 
ferous Forest, White Fir Forest, Red Fir Forest. Host Plants: LORAN- 
THACEAE: Arceuthohium (Dwarf Mistletoe), a genus occurring on 
conifers. 

55. Callophrys {Mitoura) johnsoni (Skin.) PL I, fig. 1 

JOHNSON’S HAIR-STREAK, a close relative of Callophrys {Mi- 
toura) spinetorum, may be told from the latter by means of its brownish 
upper side. The species is a prize over all of its range, but occurs more 
frequently in the Yuba Pass-Gold Lake area of Sierra County, California, 
and in the Pacific Northwest. 

Life Zones: Transition, Canadian. Plant Communities: Coniferous 
Forest. Host Plants: LORANTHACEAE: Arceuthohium Douglassii 
(Dwarf Pine Mistletoe) . 

56. Callophrys {Mitoura) nelsoni (Bdv.) PL I, fig. i 

NELSON’S HAIR-STREAK, like The California Hair-Streak, 

occurs in the nature garden behind the Yosemite Museum. Both swarm 
over the pink clusters of Spraguea { Pussy Paws ) and may be picked off 
by forceps. They may also be found at Jerseydale, at the junction of 
Highway 120 with Hetch-Hetchy Road, and on the warm road-shqulders 
above Tamarack Flat. 

Life Zones: Transition, Canadian. Plant Communities: Mixed Coni- 
ferous Forest. Host Plants: PINACEAE: Libocedrus decurrens (Incense 
Cedar) , probably. 

57. Callophrys {Incisalia) doudorojfi windi (Clench) 

PI. I, fig. j 

WIND’S ELFIN was encountered at a damp spot in the trail near 
Indian Flat, where it was flying with the following species. It differs 
from normal doudoroffi of the Monterey Coast in being much lighter 
and less heavily banded below. Males are slaty gray above and brown 
beneath; females are quite golden in color. The form has not been 
illustrated previously. 

Life Zones: Upper Sonoran. Plant Communities: Chaparral. Host 
Plants: Not known. Perhaps CRASSULACEAE: Sedum (Stonecrop), 
as for the related C. {!.) fotis. 

58. Callophrys {Incisalia) iroides (Bdv.) 

THE WESTERN ELFIN is one of the first butterflies to emerge in 
the spring. March finds it on the wing in the foothills of the San Joaquin 
Valley. It is a very plain, brown elfin. The larvae are probably bud- 
feeders, like those of Celastrina argiolus echo { Edw. ) . 


38 


GARTH AND TILDEN 


/. Res. Lepid. 


Life Zones: Upper Sonoran (lower Transition), occasionally to 
Canadian. Plant Communities: Chaparral, and the chaparral-like under- 
story of coniferous forests. Host Plants: CONVOLVULACEAE: Cuscuta 
(Dodder). RHAMNACEAE: Ceanothus (Buck Brush, Wild Lilac). 
ERICACEAE: Arhutus {M-Rdrioho) , Arctostaphylos (Manzanita), Vac- 
cinium (Bilberry). Perhaps CRASSULACEAE: Sedum (Stonecrop). 
59. Callophrys (Incisalia) eryphon (Bdv.) 

THE WESTERN BANDED ELEIN clings quite constantly to the 
coniferous forest. Never common, its discovery gives cause for comment. 

Life Zones: Transition, Canadian, lower Hudsonian. Plant Communi- 
ties: Yellow Pine, Red Eir, and Lodgepole Pine Forest. Host Plants: 
PIN ACE AE: Pinus ponder osa (Yellow Pine), P. contorta Murray ana 
(Lodgepole Pine) . 

60a. Lycaena {Tharsalea) arota arota (Bdv.) 

b. Lycaena {Tharsalea) arota virginiensis (Edw. ) 

THE AROTA COPPER and THE NEVADA COPPER occupy the 
western and eastern slopes of the Sierra Nevada, respectively. The for- 
mer was found in Pate Valley by Edmund Godwin on California Laurel, 
Umbellularia calif ornica; the latter occurs at Mono Lake and at Bodie 
where specimens seem almost indistinguishable from those from Vir- 
ginia City, Nevada, the type locality. 

Life Zones: Upper Sonoran, Transition. Plant Communities: Mixed 
Evergreen Forest (west slope); Juniper Woodland, Sagebrush Scrub 
(east slope). Host Plants: SAXIFRAGACEAE: Ribes (Currant), 
Grossularia ( Gooseberry ) . The overwintering eggs are deposited at the 
bases of thorns. 

61. Lycaena (Lycaena) gorgon (Bdv.) 

THE GORGON COPPER flies in the foothills surrounding the 
western approaches to the park. Single brooded, it is on the wing during 
late spring and early summer, when it may be expected to occur as a 
vagrant within the park boundaries. 

Life Zones: Upper Sonoran. Plant Communities: Foothill Wood- 
land, Chaparral. Host Plants: POLYGONACEAE: Eriogonum (Buck- 
wheat), especially E. nudum. 

62. Lycaena (Lycaena) heteronea Bdv. 

THE VARIED BLUE is common on the east slope of the Sierra 
Nevada. It has been found west of the divide in the sagebrush zone 
above the subalpine forest between Tioga Pass Checking Station and 
the summit of Gay lor Lakes Trail. It is also sometimes common below 
Tuolumne Meadows on the way to Return Creek, below Glen Aulin. 

Life Zones: Hudsonian (in Yosemite). Plant Communities: Sage- 
brush Scrub, often where mixed with Pine Forest or Juniper Woodland. 

Host Plants: POLYGONACEAE: Eriogonum (Buckwheat). Asso- 
ciated with E. umbellatum ( Sulphur Flower ) . 

63. Lycaena (Lycaena) xanthoides (Bdv.) 

THE GREAT COPPER flies in the foothills throughout the state 
and is distinguished by its large size and dull coloration. 


3{i):i-p6, ip6} 


YOSEMITE BUTTERFLIES 


39 


Life Zones: Upper Sonoran (lower Transition) . Plant Communities: 
Chaparral, Foothill Woodland, Yellow Pine Forest clearings. Meadow 
and open Streamside. Host Plants: POLYGONACEAE: Rumex hymen- 
osepalus (Dock). 

64. Lycaena (Lycaena) rubidus (Behr) PI. IV, fig. d 

THE RUDDY COPPER, earlier suspected of flying in the Yosemite 

region (Garth, 1935), occurs in the sagebrush region of the Gay lor 
Lakes Trail mentioned under the preceding Lycaena ( L, ) heteronea, and 
on the east Sierran slope as well. 

Life Zones: Upper Sonoran to Hudsonian (Arctic- Alpine ) . Plant 
Communities: Sagebrush Scrub, openings in Subalpine Forest, and 
Alpine Fell-Fields. Host Plants:. POLYGONACEAE: Rumex (Dock), 
probably. 

65. Lycaena (Lycaena) edit ha (Mead) 

EDITH’S COPPER is a creature of the higher elevations of the 
eastern portion of the park, where it flies in company with Lycaena 
mariposa, L. mhidus, and L. heteronea. It resembles a compact model of 
Lycaena xanthoides, and might be called ( with tongue in cheek ) , "The 
Small Great Copper.” 

Life Zones: Hudsonian, Arctic- Alpine. Plant Communities: Sub- 
alpine Forest openings, Subalpine Meadow, Alpine Fell-Fields. Host 
Plants: POLGONACEAE: Rumex paucifolius (Alpine Dock). ROS- 
ACEAE: Potentilla tenuiloba, Horkelia fusca (Cinquefoil). 

66. Lycaena (Lycaena) mariposa Keak. 

REAKIRT’S COPPER frequents small meadows of the southern 
exposure as far as timber line, but does not fly above. It was encountered 
by the senior author at 10,600 feet while crossing the Lyell Fork of the 
Tuolumne River preparatory to an ascent of Kuna Crest. 

Life Zones: Hudsonian. Plant Communities: Openings in Subalpine 
Forest. Host Plants: POLYGONACEAE: Polygonum Douglassii 
(Buckwheat); can be raised on Rumex (Dock). 

67. Lycaena (Lycaena) nivalis (Bdv. ) PL II, fig. f 

THE NIVALIS COPPER, known for many years only from a small 

colony at Glacier Point, flies generally throughout the park at elevations 
of 7,000 feet and above. It resembles The Purplish Copper, Lycaena 
(L.) helloides, but has an immaculate under side. 

Life Zones: Canadian, Hudsonian. Plant Communities: Rocky Out- 
crops and Alpine Fell-Fields. Host Plant: Unknown. POLYGON- 
ACEAE: Efiogonum (Buckwheat) suspected. 

68. Lycaena (Lycaena) helloides (Bdv.) 

THE PURPLISH COPPER is believed to be primarily a dweller of 
the lowlands, but occasional specimens have been found at higher eleva- 
tions. It lacks the luster of the preceding Lycaena (L.) nivalis and has 
a faintly dotted under side. 

Life Zones: Upper Sonoran (Transition). Plant Communities: 
Forest openings. Moist situations, rocky outcrops. Host Plants: POLY- 
GONACEAE: Rumex acetosella (Sheep Sorrel), Polygonum aviculare 
(Knotweed), Oxy theca. ONAGRACEAE: Gay ophy turn. 


40 


GARTH AND TILDEN 


/. Res. Lepid. 


69 . Lycaena (Lycaena) phlaeas hypophlaeas (Bdv.) 

PL III, fig. h 

The Sierran form of THE AMERICAN COPPER, first reported 
from the Yosemite region by Garth (1935b), has been established as 
the typical Lycaena (L.) phlaeas hypophlaeas. Although the glacial tarn 
on the north face of Mt. Maclure known locally as Bert Lake in honor 
of former park naturalist C. A. "Bert” Harwell was revisited by Oakley 
Shields, no additional specimens were found there. The north slope of 
Mt. Dana, however, yielded a limited number, found on a steep slope 
on which footing was precarious. Characterized by its small size, sub- 
dued brassy color, and bluish spots above the orange subm.arginal band 
of the secondaries, the Yosemite form appears closer to the Arctic race 
feildeni M’Lachlan than to the eastern race, to which the name americana 
Morris is now applied. 

Life Zones: Arctic- Alpine. Plant Communities: Alpine Fell-Fields 
Host Plants: POLYGONACEAE: Rumex (Dock), especially R. aceto- 
sella, for the eastern subspecies. 

70. Lycaena (Lycaena) cupreus (Edw.) PI. Ill, fig. j 

THE LUSTROUS COPPER is the most intense bit of color on the 

wing in Yosemite. Its fiery dress enlivens the mountain meadows. Only 
Lycaena (L.) snotui (Edw.) of the Rockies, considered by Klots (1936) 
and by Brown (1955) to be a subspecies of cupreus, rivals it in bril- 
liance. It flies over Spraguea umbellata (—Calyptridium umbellatum) 
(Pink Pussy Paws) on rock slides not far from retreating snow. 

Life Zones: Hudsonian, Arctic- Alpine. Plant Communities: Alpine 
Fell-Fields. Host Plants: Unknown. 

71. Lepotes marina (Reak.) 

THE MARINE BLUE, an Austral species, has been encountered by 
us only in the Mono Lake area. 

Life Zones: Lower Sonoran^ Upper Sonoran. Plant Communities: 
Several over its wide range. Host Plants: LEGUMINOSAE: Medicago 
(Alfalfa), Astragalus (Loco Weed, Rattle-Weed), Lathy rus (Sweet 
Pea). 

72. Brephidium exilis (Bdv.) 

THE PYGMY BLUE, until found at Mono Lake in the final year 
of this study, was recorded as present in the Yosemite region on the 
strength of a single female specimen obtained by Oakley Shields at Sad- 
dlebag Lake. It is the smallest of North American butterflies, with a 
wing expanse of .65 in. 

Life Zones: Unrestricted. Plant Communities: Alkali Sink, Salt 
Marsh. Host Plants: CHENOPODIACEAE: Atriplex (Salt Bush), 
Chenopodium (Lamb’s Quarters). 

73 . Hemiargus (Echinargus) isolus (Reak.) PL IV, fig. e 

REAKIRT’S BLUE is an ubiquitous species, never common, but 

occurring over a wide territory and to a considerable elevation in the 


2( i ): i ~96, 1963 


YOSEMITE BUTTERFLIES 


41 



Fig. 6. Moist meadow surrounded by Lodgepole Pine and Red Fir 
forest. Canadian Zone. Research Reserve, el. 8,000 ft. Anthocharis sara 
Stella, Plejus saepiolus, Polygonia zephyrus, Precis orythya evarete, and Pyrgus 
ruralis may be seen here.^ — ^Neva Snell. 


42 


GARTH AND TILDEN 


/. Res. Lepid. 


drier eastern portions of the Sierra Nevada, at least in the 1958 season. 
A row of black dots encircled with white on the underside of the pri- 
maries is a distinguishing feature. 

Life Zones: Unrestricted. Plant Communities: (Among others) 
Sagebrush Scrub. Host Plants: LEGUMINOSAE: Astragalus (Rattle 
Weed, Loco Weed), Hosackia ( = Lotus), Trifolium (Clover). 

74. Lycaeides ar gyro gnomon anna (Edw. ) PL II, fig. i 

THE ANNA BLUE occurs at higher elevations in the western por- 
tion of the park, being found in the upper tier of timbered zones. It 
is always associated with lupine. 

Life Zones: Canadian, Hudsonian. Plant Communities: Fir Forest 
openings. Host Plants: Undetermined. Probably LEGUMINOSAE: 
Lupinus (Lupine). 

75. Lycaeides melissa inyoensis Nab. PL IV, figs, i and 1 

THE INYO BLUE is the name to be given to the race formerly 

recognized by most Californian lepidopterists as lotis ( Lint. ) . According 
to Nabokov (1949), true lotis is a race of the foregoing Lycaeides 
ar gyro gnomon restricted to peat bogs of Mendocino County. THE INYO 
BLUE flies in the eastern portion of the Yosemite region. 

Life Zones: Unrestricted. Plant Communities: Several, including 
Freshwater Marsh, Alkali Sink, Meadowland, Marsh, Cultivated Field, 
Fence-Row, and Roadside Associations. Host Plants: LEGUMINOSAE: 
Astragalus (Loco Weed, Rattle Weed), Hosackia { — Lotus), Lupinus 
(Lupine), Medicago (Alfalfa), Glycyrrhiza lepidota (Liquorice), at 
Lone Pine. 

76. Plebejus {Agriades) glandon podarce (F. & F.) 

PL III, fig. g 

THE GRAY BLUE is perhaps the most characteristic butterfly of 
the Sierran sub-alpine meadow. It is common at Tioga Pass, where at 
times it fairly swarms, always in association with Dodecatheon alpinum 
(Shooting Star). As soon as the sun’s rays leave the mountain valleys, 
it settles upon the grasses and sedges and may be picked by hand or by 
forceps. A cross on each fore-wing serves to distinguish the species. 

Life Zones: Upper Canadian, Hudsonian (Arctic- Alpine ) . Plant 
Communities: Sub-alpine Meadow, Alpine Fell-Fields. Host Plants: 
Unknown. PRIMULACEAE: Gregoria for European P. glandon, to 
which Primula (Primrose) and Dodecatheon (Shooting Star) are re- 
lated. 

77. Plebejus {Plebejus) saepiolus (Bdv.) 

THE GREENISH BLUE flies with the foregoing species, Plebejus 
(A.) glandon podarce, and in equal abundance. P. (P.) saepiolus, how- 
ever, descends to lower elevations, as at Glacier Point and Aspen Valley. 
As with the former species, only the male wears the color indicated by 
the common name, the female being drab brown. The commonest blue 
in the park at higher elevations, and the one with the longest flight 
season, it has two broods, the first tending to oviposit on clover, the 
second on Hosackia. 


2(i);i-96, 1963 


YOSEMITE BUTTERFLIES 


43 


Life Zones: Canadian, Hudsonian (Arctic- Alpine). Plant Com- 
munities: Subalpine Meadow, Alpine Fell-Fields, Fiost Plants: LEGUM- 
INOSAE: Trifolium (Clover), Hosackia (= Lotus). 

78a. Pie be jus {Icaricia) icarioides icarioides (Bdv.) 
b. Plebejus {Icaricia) icarioides helios (Edw. ) 

BOISDUVAL’S BLUE is a widely distributed species, having been 
subdivided into numerous geographical races. The Yosemite form is 
close to the nominate form; the name helios Edw. is here applied to spec- 
imens from Mono Lake. Because of its relatively large size and uniform 
markings, as well as its solid blue coloration, this species may be confused 
with Lycaena (L.) heteronea Bdv. when on the wing. 

Life Zones: (Upper Sonoran), Transition, Canadian. Plant Com- 
munities: Coastal Sage Scrub, Chaparral, Foothill Woodland, Yellow 
Pine and Mixed Coniferous Forest openings for P. i. icarioides; Sage- 
brush Scrub, Pinon-Juniper Woodland, Juniper Woodland, Jeffrey Pine 
Forest openings for P. i. helios. Flost Plants: LEGUMINOSAE: Lfipinus 
(Lupine) , perennial species only. 

79 . Plebejus (Icaricia) shasta comstocki (Fox) PL III, fig. e 

THE YOSEMITE BLUE is the name given this insect in Comstock’s 

"Butterflies of California,” although by virtue of its Latin name it might 
be called Comstock’s Blue instead. Its metropolis is above timber line, 
but isolated colonies occur above 7,000 feet, as at Glacier Point. It flies 
over dwarf lupine and yellow buckwheat, Eriogonum incanum, but has 
not been observed to oviposit on either. The long flight season suggests 
two broods. The Rocky Mountain Plebejus (Icaricia) shasta minnehaha 
(Scud.) flies only above tree line and often above 12,000 feet. 

Life Zones: Hudsonian, Arctic- Alpine. Plant Communities: Sub- 
alpine Forest openings, Alpine Fell-Fields. Host Plants: Unknown. LE- 
GUMINOSAE: Lupinus (Alpine Lupine) suspected. 

80. Plebejus ( Icaricia ) acmon ( West. & Hew. ) 

THE ACMON BLUE may be found throughout the Sierra Nevada 
at almost any elevation. It occasionally flies in company with the follow- 
ing Plebejus (I.) lupini. From this circumstance, plus a possible differ- 
ent host plant, it has been suggested that the two are distinct species, and 
they are tentatively so considered here. 

Life Zones: Upper Sonoran, Transition, Canadian (Hudsonian. Plant 
Communities: Many. Host Plants: LEGUMINOSAE: Astragalus (Loco 
Weed, Rattle Weed), Hosackia ( — Lotus) , Lupinus (Lupine) . POLY- 
GON ACE AE : Eriogonum ( Buckwheat ) . 

81. Plebejus (Icaricia) lupini (Bdv.) 

THE LUPINE BLUE may be distinguished from Plebejus (I.) 
acmon by the purplish luster of both sexes. The two are sometimes 
found together, as at Smoky Jack Camp. Specimens from 11,000 feet 
are visibly smaller than those from moderate elevations; those from Lee 
Vining are visibly larger. 

Life Zones: Canadian, Hudsonian, High Arid Transition. Plant 
Communities: Not determined. Subalpine Forest, Alpine Fell-Fields, 


44 


GARTH AND TILDEN 


/. Res. Lcpid. 


Juniper Woodland, Sagebrush Scrub, among others. Host Plants: Ap- 
parently unknown, as distinct from P. (I.) acmon. Eriogonum (Buck- 
wheat) association noted. 

82. Everes comyntas (Godt.) 

THE EASTERN TAILED BLUE is the species represented on the 
east side of the Sierra Nevada, although it is not known to which of 
several subspecies the Mono Lake population should be assigned. 

Life Zones: Upper Sonoran (Hudsonian). Plant Communities: Not 
determined. Freshwater Marsh, Moist Meadow associations. Host Plants: 
LEGUMINOSAE: Trifolium (Clover), Vicia (Vetch), Astragalus 
(Loco Weed, Rattle Weed). 

83. Everes amyntula 

THE WESTERN TAILED BLUE has been seen in Yosemite Park 
by one of us ( Garth ) . It was flying over a tall, succulent lupine in the 
vicinity of a white fir grove at Hetch-Hetchy summit. It occurs in the 
foothills west of the park. 

Life Zones: Transition (Canadian). Plant Communities: Yellow 
Pine Forest openings. Host Plants: LEGUMINOSAE: Astragalus (Loco 
Weed, Rattle Weed), Lathyrus (Sweet Pea), Vicia (Vetch). 

84. Philotes enoptes ( Bdv. ) 

THE DOTTED BLUE, as found at Glacier Point, is typical. Speci- 
mens found at Research Reserve show mixed characters, but are separable 
from the following species. 

Life Zones: Transition, Canadian (Hudsonian). Plant Communities: 
Understory of several. Host Plants: POLYGONACEAE: Eriogonum 
(Buckwheat) of the latifolium group. The eggs are concealed in the 
flower head, on which the larvae feed. 

85a. Philotes hattoides hattoides (Behr) 
b. Philotes hattoides glaucon ( Edw. ) 

THE SQUARE-SPOTTED BLUE of Yosemite is an inhabitant of 
the Alpine Fell-Fields, appearing very early in the year for so high an 
altitude. It also flies atop Research Ridge (Boundary Hill), keeping 
well within the few acres marked by the presence of weak Hudsonian 
elements. It flies over Eriogonum incanum, a yellow buckwheat, and 
Astragalus bolanderi, a straggly legume. Specimens found within the 
park represent the typical form. Specimens found at Mono Lake are 
referred to THE GLAUCON BLUE, a subspecies. 

Life Zones: Hudsonian for P. h. hattoides; Upper Sonoran, Arid 
Transition for P. b. glaucon. Plant Communities: Rock Garden for P. h. 
hattoides; Juniper Woodland, Sagebrush Scrub for P. b. glaucon. Host 
Plants: POLYGONACEAE: Eriogonum (Buckwheat). 

86. Philotes speciosa ( Hy. Edw. ) 

THE SMALL BLUE is typically a desert insect, found around dry 
lake beds (playas) in the Mojave Desert. Occasional specimens occur 
in the Central Valley region of California, and one of us (Garth) has 
found it at Hmne Lake in Sequoia-Kings Canyon National Park. The 


2(t):i-96, 1963 


YOSEMITE BUTTERFLIES 


45 


Mariposa and Briceburg specimens collected by G. and R. Bohart may 
represent an undescribed subspecies. Both specimens are larger, with 
wider margins in the male, and with the hind wings less spotted below, 
than normal specimens from the desert. In the absence of sufficient 
material, description must wait. 

Life Zones: Lower Sonoran in the Mojave Desert; Upper Sonoran 
(Transition) in the western Sierra Nevada. Plant Communities: Not 
determined. Stream Bed Association. Host Plants: POLYGONACEAE: 
Eriogonum (Buckwheat) in San Diego County; Oxy theca perfoliata in 
the Mojave Desert; in the Sierra unknown. 

87» Phaedrotes piasus i'B&v.) 

THE ARROWHEAD BLUE, while not common in the Yosemite 
region, occurs in the western foothills, the middle elevations, and on the 
eastern slope at Mono Lake. 

Life Zones: Upper Sonoran, Transition, Canadian. Plant Commun- 
ities: Several. Host Plants: LEGUMINOSAE: Lupinus (Lupine). 

88a. Glaucopsyche lygdamus hehrii ( Edw^. ) 
b. Glaucopsyche lygdamus Columbia (Skin.) 

BEHR’S BLUE flies in the early spring in the western foothills. 
At higher elevations it is replaced by THE COLUMBIA BLUE, a 
montane subspecies found in cooler parts of most of the United States 
and Canada. It is not common in the Tioga Pass region, nor indeed 
anywhere, its colonies being widely scattered. 

Life Zones: Upper Sonoran for G. 1. hehrii; Canadian, Hudsonian 
for G. 1. Columbia. Plant Communities: Foothill Woodland, Chaparral 
for G. 1. hehrii; Subalpine Forest, Alpine Fell-Fields for G. /. Columbia. 
Host Plants: LEGUMINOSAE: Lottis (= H os ac kia) , Astragalus (Loco 
Weed, Rattle Weed), Lathyrus (Sweet Pea), Lupinus (Lupine), Vida 
(Vetch). 

89. Celastrina argiolus echo (Edv^. ) 

THE ECHO BLUE is the western representative of a highly poly- 
morphic species, now recognized as having Old World affinities. One 
of the earliest butterflies to emerge in the spring, it follows the season’s 
advance to higher altitudes, occurring through three life zones. Male 
and female are strikingly dissimilar in pattern. 

Life Zones: Upper Sonoran, Transition, Canadian. Plant Commun- 
ities: Many. Host Plants: LEGUMINOSAE: Hosackia (= Lotus), 
Lupinus (Lupine); RHAMNACEAE: Ceanothus (Lilac), Spiraea 
(Spirea); SAPINDACEAE: Aesculus (Buckeye); ERICACEAE: 
Arctostaphylos (Manzanita); FAGACEAE: Quercus (Oak); CORN- 
ACEAE: Cornus (Dogwood). In many instances, only the flower buds 
are fed upon. 



PLATE I 


3( i ): i -96, 1963 


YOSEMITE BUTTERFLIES 


47 


PLATE I 

THE WESTERN FOOTHILLS 

a. Thorbes diversus 

Aspen Valley, J.S.G. 

b. Argynnis z. zerene cf 

Mather, J.S.G. 

c. Coenonympha t. calif ornica cf 

Jerseydale, J.S.G. 

d. Euphydryas e. rubicunda d 

Indian Flat, J.W.T. 

e. Ce rcyonis sylvestris 9 

Hetch-Hetchy, J.S.G. 

f. Anthocharis lanceolata d 

Yosemite Valley, Paul’ Allen 

g. Limenitis b. californica 9 

Hetch-Hetchy, J.S.G. 

h. Habrodais grunus 9 

Hetch-Hetchy, J.S.G. 

i. Callophrys nelsoni 9 

Hetch-Hetchy, J. S.G. 

j. Callophrys d. windi 9 

Indian Flat, J.W. t. 

k. Strymon auretorum 9 

Hetch-Hetchy, J S.G. 

l. Callophrys johnsoni d 

Jerseydale, A.O.S. 

PLATE m 

THE HIGH COUNTRY 

a. Melitaea d. malcolmi d 

Gaylor Lakes, J.S.G.. 

b. Oeneis c. ivallda d 

Glacier Lodge, J.S.G. 

c. Euphydryas e. nubigena 9 

Rafferty Creek, J.S.G. 

d. Polites s. tecunaseh 

Upper Gaylor Lake, J.S.G. 

e. Plebejus s. comstocki d 

Yosemite Park, J.S.G. 

f. Parnass|us p. behri d 

Gayl Lakes, J.S.G. 

g. Plebejus g. podarce d 

Research Reserve, J.S.G. 

h. Lycaena p. hypophlaeas 9 

N. slope, Mt. Dana, A.O.S. 

i. Colias behri d 

Tioga Pass, J.S.G. 

j. Lycaena cupreus d 

Lembert Dome, J.S.G. 

k. Callophrys lemberti (paratype) d 

Tioga Pass, J.W.T. 

l. Argynnis m. arge 9 

Tioga Pass, J.S.G. 

m. Hesperia miriamae 9 

Unicorn Peak, A.O.S. 


PLATE II 

THE WESTERN MID- ELEVATIONS 

a. Anthocharis s. Stella 9 

White Wolf, J.S.G. 

b. Limenitis lorquini d 

Aspen Valley, J.S.G. 

c. Phyciodes mylitta 9 

YosemiLe Creek, J.S.G. 

d. Boloria epithore 9 

Tioga Road, J.S.G. 

e. Parnassius c. baldur d 

White Wolf, J.S.G. 

f. Lycaena nivalis 9 

Research Reserve, J.S.G. 

g. Hesperia harpalus cT 

Yosemite Creek, J.S.G. 

h. Argynnis a. irene rf 

Aspen Valley, J.S.G. 

i. Lycaeides a. anna cf 

Aspen Valley, J.S.G. 

j. Melitaea palla d 

White Wolf, J.S.G. 

k. Polygonia zephyrus 9 

Tuolumne Grove, J.S.G. 

l. Melitaea hoffmanni 9 

White Wolf, J.S.G. 

PLATE IV 

THE MONO BASIN 

a. Coenonympha t. mon o d 

W of LeeVining, A.O.S. 

b. Argynnis c. nevadensis d 

near Mammoth, J.S.G. 

c. Cercyonis oeta d 

June Lake, J.S.G. 

d. Lycaena rubidus d 

Virginia Lakes, J.S.G. 

e. Hemiargus isolus 9 

Rafferty Cr., J.S.G. 

f. Euphydryas e. monoensis d 

Mono Lake, A.O.S. 

g. Strymon behrii 9 

Mammoth, J.S.G. 

h. Limenitis w. nevadae d 

Mono Lake, J.S.G. 

i. Lycaeides m. inyoensis ff 

Mono Lake, J.W.T. 

j. Strymon f. semiluna 

Mono Lake, A.O.S. 

k. Argynnis n. apacheana (f 

Round Valley, J.S.G. 

l. Lycaeides m. inyoensis 9 

Lone Pine, J.W.T. 



PLATE II 




PLATE III 


50 


GARTH AND TILDEN 


/. Res. Lepid. 


PIERIDAE 

90a, Anthocharis {Anthocharis) sara saral^wc^is 

Anthocharis {Anthocharis) sara reakirtii Edw., gen. vern. 

b. Anthocharis {Anthocharis) sara Stella Edw. PI. II, fig. a 

THE SARA ORANGE-TIP and its overwintering form, REA- 
KIRT’S ORANGE-TIP, occur in the western lowlands at Briceburg and 
El Portal. THE STELLAR ORANGE-TIP is the high-altitude race 
found in California, race jzdia Edw. being from Colorado and race flora 
Wgt. from Oregon and north. Females of this form are yellowed, un- 
like those of the lowland forms, in which females are white like the 
males. 

Life Zones: Upper Sonoran, lower Transition for A. sara sara; upper 
Transition, Canadian for A. s. Stella. Plant Communities: Valley Grass- 
lands, Foothill Woodland, Chaparral, Yellow Pine Forest for A. s. sara; 
White Fir Forest, Red Fir Forest for A. s. Stella. Forest openings and 
Riparian Association. Host Plants: CRUCIFER AE: Arabis (Rock 
Cress), Brassica (Mustard), Sisymbrium (Hedge Mustard). 

91. Anthocharis {Falcapica) lanceolata Lucas PI. I, fig. f 

THE LANCEOLATE MARBLE (formerly called Boisduvals 

Marble when authorship was attributed to him) is an early flier, to be 
sought from mid-May to mid- June. It seeks the vertical walls of Yose- 
mite Valley and may be seen on the precipitous talus slopes near Bridal 
Veil Falls. Yosemite specimens are almost gray enough beneath to match 
race australis { Grin. ) from southern California. 

Life Zones: Upper Sonoran, Transition. Plant Communities: Chap- 
arral, Foothill Woodland, Mixed Evergreen Forest. Rocky Outcrop and 
Cliffside Associations. Host Plants: CRUCIFER AE, especially Arabis 
perfoliata (Rock Cress). 

92. Euchloe creusa hy antis (Edw.) 

EDWARDS’ MARBLE may be separated from the following Colo- 
rado Marble by the heavy green marbling of the under side of the 
secondaries against a background of dead white, Euchloe ausonides 
coloradensis having less green and of a decidedly yellowish cast against 
a pearly white ground color. The green of both species is an effect 
produced by yellow scales laid over black, as examination with a hand 
lens will demonstrate. The black bar at the end of the cell is wide in 
creusa. 

Life Zones: Upper Sonoran, Transition, Canadian. Plant Commun- 
ities: Chaparral, Foothill Woodland, Mixed Evergreen Forest, White Fir 
Forest. Host Plants: CRUCIFERAE: Arabis (Rock Cress), Streptan- 
thus (Jewel Flower). 

93. F.uchloe ausonides coloradensis (Hy. Edw.) 

THE COLORADO MARBLE is the common Euchloe of the Re- 
search Reserve, where approximately four specimens were found for 
every one of E. creusa hyantis. Both fly to the mountain tops and may 
be seen in good numbers by the observer who stations himself in such 



PLATE IV 


52 


GARTH AND TILDEN 


/. Res. Lepid. 


a situation. Like Behr’s Parnassian, Parnassius phoebus behrii Edw., it 
belongs to the Great Basin fauna. The black bar at the end of the cell 
is narrow. 

Life Zones: Canadian, Hudsonian. Plant Communities: Coniferous 
Forest openings. Host Plants: CRUCIFER AE: Arabis (Rock Cress), 
Erysimum (Wall Flower), Sisymbrium (Hedge Mustard). 

94. CoUas eury theme Bdv. 

THE CLOUDED SULPHUR is partial to grassy meadows and to 
cultivated areas, where it substitutes introduced alfalfa for indigenous 
Astragalus for its food plant. The summer form, amphidusa Bdv., is 
larger and more orange. Both have two forms of the female, normal 
and albinic, but only One form of the male. 

Life Zones: Unrestricted. Plant Communities: Immaterial, if 
legumes be present. Host Plants: LEGUMINOSAE: Astragalus (Loco 
Weed, Rattle Weed), Hosackia (— Lotus) ^ Medicago (Alfalfa), Mel- 
ilotus (Sweet Clover), Trifolium (Clover), Vida (Vetch). 

95. CoUas philodice hagenii Edw. 

HAGEN’S SULPHUR is a Great Basin race of a widely ranging 
North American species found otherwise mostly east of the Rocky 
Mountains. According to Hovanitz (1943, 1950), it does not occur 
west of the Sierra-Cascade Divide Triple-brooded at Mono Lake, it 
may be recognized by its yellow, rather than orange, coloration. 

Life Zones: Upper Sonoran, Transition. Plant Communities: Moist 
Meadows. Host Plants: LEGUMINOSAE: Trifolium (Clover). 

96. CoUas occidentaUs chrysomelas Hy. Edw. 

THE GOLDEN SULPHUR is the largest member of its genus 
occurring in California. An early flier, it is considered a rarity, having 
been seen by but one of us (Tilden) in the vicinity of Mather on June 
30, 1962, in the course of this survey. 

Life Zones: Transition. Plant Communities: Associated with con- 
iferous forest at moderate elevations. Host Plants: LEGUMINOSAE: 

( Legumes ) . 

97. CoUas behrii Edw. PI. Ill, fig. i 

BEHR’S SULPHUR is the only greenish sulphur butterfly of the 

Sierra Nevada. Until the opening of the Tioga Road in 1915 it was 
considered a great rarity because of the extreme inaccessibility of its 
habitat, the Tuolumne Meadows. John Batiste Lembert, who homestead- 
ed the Soda Springs quarter section in 1885, found that there was a 
demand for these butterflies and for a dozen years supplied them in 
quantity to universities and museums. The secret of their haunts he 
shared with the Indians, who in turn refused to divulge it to outsiders. 

We now know that CoUas behrii has as its metropolis the Tuolumne 
Meadows, that smaller colonies are present in adjacent subalpine 
meadows, and that it has been found to the south at Rock Creek and 
Mineral King. Its occurrence is tied closely to the distribution of 
Dwarf Bilberry, on which the larvae exclusively feed. Form 9 canescens 


i$6s 


YOSEMITE BUTTERFLIES 


53 


PLATE V 

a. Erynnis pacuvius lilias d 

Chowchilla Mtn. J.W.T. 

b. Argynnis zerene malcolmi d 

Mammoth Lakes, J.W. Friday 

c. Lycaena gorgon 9 

Lebec, J.S.G. 

d. Everes amyntula d 

Yucaipa, J.S.G. 

e. Papilio indra d 

Tioga Pass, A.O.S. 

f. Callophrys iroides d 

Grant Nat'l Park J.S.G. 

g. Strymon saepium 9 

Hetch-Hetchy, J.S.G. 

h. Lycaena editha 9 

Tioga Pass, J.S.G. 

i. Philotes battoides glaucon d 

Mono Lake. A.O.S. 

j. Brephidium exilis 

Long Beach, J.S.G. 

k. Philotes battoides battoides d 

Research Reserve, J.S.G. 

l. Pieris occidentalis calyce 9 

W of Tioga Pass, A.O.S. 

m. Danaus plexippus d 

Mather, J.S.G. 

n. Euchloe creusa lotta 9 

Darwin, R.P. Allen 

o. Polites sonora 9 

Aspen Valley, J.S.G. 

p. Vanessa carye d 

Topanga Canyon, J.S.G. 

q. Vanessa cardui 9 

Grant Nat'l Park, J.S.G. 

r. Polites sonora d 

Bridal Veil Creek, J.S.G. 

s. Lycaena heteronea 9 

Gem Lake, J.S.G. 

t. Callophrys iroides d 

Mammoth Lakes, J.S.G. 

u. Phaedrotes piasus d 

Tujunga, J.S.G. 

V. Plebejus icarioides helios 9 
Warren Creek, J.S.G. 


PLATE VI 

a . Thorbes pylades j 

Jersydale, J.W.T. 

b. Pyrgus ruralis cf 

Boulder Creek, J.W.T. 

c. Ochlodes agricola d 

Hetch-Hetchy, J.S.G. 

d. Amblyscirtes vialis , d 

Flagstaff, Ariz., K. Roever 

e. Argynnis atlantis irene d 

Aspen Valley, J.S.G, 

f. Leptotes marina 9 

Long Beach, J.S.G. 

g. Vanessa virginiensis 9 

Santa CruzMtns., J.S.G, 

h. Strymdn melinus pudica d 

Long Beach, J.S.G. 

i. Nymphalis milberti 9 

Yucaipa, J.S.G. 

j. Lycaena arota virginiensis 9 

Mono Lake, A.O.S. 

k. Pieris napi venosa d 

Briceburg, K. T. 

l. Papilio eurymedon ^ 

Idyllwild, J.S.G. 

m. Plebejus saepiolus (f 

Research Reserve, J.S.G. 

n. Colias eurytheme cf 

Mather, J.S.G. 

o. Argynnis cybele leto 9 

Strawberry, J.S.G. 

p. Argynnis callippe inornata 9 

Jerseydale, A.O.S. 


54 


GARTH AND TILDEN 


Lepid. 


J. Res. 



PLATE V 



YOSEMITE BUTTERFLIES 


55 


s(i):i-$6) 1963 






56 


GARTH AND TILDEN 


/. Res. Lepid. 


Comst., although relegated to synonymy by McDunnough (1938), we 
find to be a valid albinic form that may be recognized on the wing. 

Life Zones: Hudsonian (Arctic- Alpine ) . Plant Communities: 
Subalpine Meadows; basin areas of Subalpine Forest; moister parts of 
Alpine Fell-Fields. Host Plants: ERICACEAE: Vaccinium caespitosum 
(Dwarf Bilberry). 

98. Colias {Zerene) eurydice^dY. 

THE CALIFORNIA DOG-FACE or FLYING PANSY, as it is 
sometimes called, must be a vagrant in the Yosemite region, for Hall 
(1912) does not record its food plant, False Indigo or Lead Bush, as 
occurring in the park. It is abundant in the foothills of southern Cali- 
fornia, and has been selected as the State butterfly. The male bears the 
figure of a dogs head on the forewing; the female is pure sulphur 
yellow. 

Life Zones: Upper Sonoran, lower Transition. Plant Communities: 
Not determined for the Yosemite region. Host Plants: LEGUMI- 
NOSAE: Amorpha calif ornica (False Indigo). 

99. Nathalis iole Bdv. 

THE DAINTY YELLOW is likely to be encountered in the 
least expected places, from sea level to 9,000 feet, and from early 
spring to early winter. There are no set rules of behavior for this 
diminutive sulphur, which has followed the Filaree into many an 
improbable situation. 

Life Zones: Unrestricted. Plant Communities: Ubiquitous. Host 
Plants: GERANIACEAE: Erodmm (Filaree) . COMPOSITAE: Helen- 
ium ( Sneezeweed ) , Dyssodia (None). CARYOPHYLLACEAE: Stel- 
laria ( Chickweed ) . 

100. Neophasia menapia tau (Scud.) 

THE PINE WHITE reaches the peak of its flight in August. 
The adults are high fliers, seldom descending from the tree tops. Al- 
though not abundant in 1933, they were seen by the hundreds in 
1928 along the Big Oak Flat road by one of us (Garth) . The larvae are 
of special interest to forest entomologists, having been known to 
defoliate pine trees over large areas. Resurrected from synonymy, the 
name tau is now applied to the western form. 

Life Zones: Transition, Canadian. Plant Communities: Yellow 
Pine Forest, Jeffrey Pine Forest (east slope); occasionally Lodgepole 
Pine Forest. Host Plants: PINACEAE: Pinus ponder osa (Yellow 
Pine), P. contorta Murray ana (Lodgepole Pine); also, according to 
the observation of one of us (Tilden), Pseudotsuga taxifolia (Douglas 

101. Pieris (Pontia) beckeriiEdw. 

BECKER’S WHITE prefers an arid environment and is encountered 
east of the Sierra Nevada from the Mojave Desert north through 
Owens Valley to Mono Lake. The green-margined veins of the under 
side suggest one of the Marbles, and where it flies with Anthocharis 




YOSEMITE BUTTERFLIES 


57 


PLATE Vn 

a. Epargyreus clarus 9 

Lagunitas, E.C. VanDyke 

b. Hesperia juba tf 

Mather, J.W.T. 

c. Ochlodes sylvanoides d* 

Virginia Lakes, J.S.G. 

d. Erynnis propertius d 

Yosemite Creek, J.S.G. 

e. Glaucopsyche lygdamus Columbia d 

White Wolf, J.S.G. 

f. Cercyonis pegala ariane 9 

Round Valley, J.S.G. 

g. Argynnis egleis d 

Yosemite Creek, J.S.G. 

h. Plebejus i. icarioides 9 

Aspen Valley, J.S.G. 

i. Euchloe creusa lotta 9 

Mono Lake, A.O.S. 

j. Philotes speciosa d 

Mojave Desert, J.S.G. 

k. Heliopetes ericetorum d 

Switzer-land, J.S.G. 

l. Apodemia m. mormo 

Mono Lake, A.O.S. 

m. Papilio rutulus d 

Idyllwild, J.S.G, 

n. Precis orithya evarete d 

Grant Nat'l Park, J.S.G, 

o. Plebejus lupini 

W, of Tioga Pass, A.O.S. 

p. Nymphalis californica 9 

Idyllwild, J.S.G. 

q. Colias eurytheme form alba 9 

Big Bear Lake, J.S.G. 

r. Lycaenopsis argiolus echo 9 

Yucaipa, J.S.G. • 

s. Lycaena a. arota 9 

Gold Creek, J.W.T. 

t. Lycaena xanthoides 9 

Mather, J.S.G. 

u. Callophrys spinetorum d 

Idyllwild, J.S.G. 

V. Strymon adenostematis 9 
Hetch-Hetchy, J.S.G. 


PLATE Vm 

a* Erynnis tristis d 

Redwood City, J..W.T. 

b. Thorbes mexicana nevada d 

Tioga Pass, A.O.S. 

c. Pyrgus communis d 

Los Angeles, J.S.G. 

d. Nathalis iole d 

Warren Creek, A.O.S. 

e. Pieris sisymbrii d 

Research Reserve, J.S.G. 

f. Ochlodes agricola d 

El Portal 

g. Strymon dr yop e (f 

Mono Lake. A.O.S. 

h. Papilio zelicaon 9 

Mather, J.S.G. 

i. Strymon californica d 

Hetch-Hetchy 

j. Lycaena helloides d 

Mather, J.S.G. 

k. Lycaena nivalis 9 

Glacier Point, J.D. Gunder 

l. Melitaea leanira d 

m. Colias (Zerene) eurydiced 

Santa AnaR,, J.S.G, 

n. Phyciodes c. campestris d 

Mather , J. S. G. 

o. Argynnis mormonia arge d 

Hetch-Hetchy, J.S.G. 

p. Plebejus acmon d 

Hatch- Hetchy 

q. Philotes enoptes 9 

Research Reserve, J.S.G. 

r. Polygonia satyrus d 

Mono Lake, A.O.S, 


/. Re$. Lepid. 


58 


GARTH AND TILDEN 

PLATE VII 



V 


4 . 


2(i):i-96, 196} 


YOSEMITE BUTTERFLIES 


59 



PLATE 












60 


GARTH AND TILDEN 


/. Res. Lepid. 


lanceolata australis Grin, in some of the desert canyons the two are 
not always distinguishable when on the wing. 

Life Zones: Upper Sonoran, Transition. Plant Communities: Sage- 
brush Scrub, Shadscale Scrub, Pihon-Juniper and Juniper Woodland. 
Host Plants: CAPPARIDACEAE: Isomeris arbor ea (Bladder Pod). 
CRUCIFER AE: Brassica nigra (Mustard), Sisymbrium altissimum 
( Hedge Mustard ) , Stanley a pinnata ( Desert Plume, Prince s Plume ) . 

102. Pieris {Pontia) sisymbriiBd\. 

THE CALIFORNIA WHITE is an early flier in the western 
foothills, where it may be found at Briceburg and El Portal in mid- 
March. At higher elevations it flies in mid-summer with the follow- 
ing Pieris {Pontia) occidentalis calyce, these two being the only whites 
then on the wing. While Sisymbrium, from which it takes its scientific 
name, grows abundantly in the meadows of Research Reserve, the 
species exhibits a marked preference for Caulanthus and Streptanthus 
when these are available, according to Hovanitz (personal communica- 
tion). 

Life Zones: Unlimited. Plant Communities: The entire gamut from 
west to east. Host Plants: CRUCIFER AE: Arabis (Rock Cress), 
Caulanthus (Rock Cabbage), Sisymbrium (Hedge Mustard), Strep- 
tanthus (Jewel Flower). 

103. Pieris {Pontia) protodice Bdv. & Lee. 

THE COMMON WHITE appears limited to the lower tier of 
life zones in Yosemite. Those seen above Transition Zone are believed 
to have wandered upward because of their worn condition. 

Life Zones: Upper Sonoran, Transition (Canadian). Plant Com- 
munities: Many. Host Plants: CAPPARIDACEAE: Cleome lutea (Bee 
Plant). CRUCIFERAE: Brassica nigra (Mustrad), Raphanus (Rad- 
ish), Lepidi'um densiflorum (Pepper Grass), Sisymbrium altissimum 
(Hedge Mustard), the first and two last at Mono Lake on the authority 
of W. Hovanitz. 

104a. Pieris {Pontia) occidentalis occidentalis Reak. 

b. Pieris {Pontia) occidentalis calyce Edw^. 

THE WESTERN WHITE occurs throughout the park, its altim- 
dinal form, EDWARD’S WHITE, at higher elevations only, although 
a small, dark form resembling P. o. calyce may be found in early 
spring in the western foothills. What one finds in the high country 
in August is calyce, freshly emerged (It is the only brood at higher 
elevations), plus worn second-brood P. o. occidentalis that have in- 
vaded from lower levels. L. G. Higgins of England recently (1953) 
associated calyce Edw. with the European callidice Esper as subspecies 
of P. {Pontia) occidenalis, which has been shown by Hovanitz (1963) 
to be specifically distinct from the preceding P, {Pontia) protodice. 

Life Zones: Unrestricted for P. o. occidentalis; Canadian, Hudsonian 
for P. 0 . calyce. Plant Communities: Many for P, o. occidentalis; Alpine 




YOSEMITE BUTTERFLIES 


61 


Fell-Fieids for P. o. calyce. Host Plants: CRUCIFERAE: Arabis (Rock 
Cress), Lepidium densijlorum (Pepper Grass), Sisymbrium altissimum 
(Hedge Mustard), the last two in the Mono Lake area, according to 
Hovanitz (personal communication). 

105. Pieris {Pieris) napi venosa Scud, 

THE VEINED WHITE is found in the foothills adjacent to Yose- 
mite National Park on the San Joaquin Valley side from mid-March 
to mid-May, when it appears on the floor of Yosemite Valley. More 
lightly marked than typical venosa, this foothill population is not 
referable to any other named subspecies. 

Life Zones: Upper Sonoran, Transition. Plant Communities: Foot- 
hill Woodland, Chaparral, Mixed Coniferous Forest. Riparian and 
Cool Woodland associations. Host Plants: CRUCIFERAE: Barb area 
(Winter Cress), Brassica (Mustard), Dentaria (Milk Maids), Rapha- 
nus ( Radish ) . 

106. Pieris (Pieris) rapae (Linn.) 

THE CABBAGE WHITE was introduced from Europe into Que- 
bec, Canada, in 1858, reaching California about 1883. It now occupies* 
not only cultivated areas, but has expanded into wild lands as well. 

Life Zones: Unrestricted. Plant Communities: Many, as would be 
expected of a highly adaptable immigrant. Host Plants: CRUCIFERAE: 
Brassica (Mustard), Radicula (Yellow Cress), the latter around 
Cathay. 

PAPILIONIDAE 

107. Papilio zelicaon Luc. 

THE ANISE SWALLOWTAIL is a "hill-topper” species, choosing 
the most exposed situations for its flight. Usually only the males disport 
themselves in this way, six being found at the very summit of Research 
Ridge. It is partial to the blossoms of the Western Wallflower, Erysi- 
mum asperum. Although readily confused with Papilio bairdii brucei 
Edw., which occurs in the region of the Mono Craters east of the Sierra 
Nevada, P. zelicaon has less yellow on the abdomen than does P. brucei. 

Life Zones: Unrestricted. Plant Communities: Chaparral, Foothill 
Woodland, openings in Coniferous Forest. Host Plants: UMBEL- 
LIFER AE: Daucus (Wild Carrot), Poeniculum vulgar e (Fennel, An- 
ise), Heracleum lanatum (Cow Parsnip). 

108. Papilio indr a Reak. 

THE INDRA SWALLOWTAIL has been seen also at the summit 
of Research Ridge. An erratic flier, it has proven most elusive, and 
Yosemite records are few. These include the floor of Yosemite Valley, 
the summit of Yosemite Falls, northwest above Tioga Pass, and Mono 
Lake. On the west bank of Yosemite Creek it flies with Parnassius 
phoebus behrii Edw. on dry slopes in association with Juniper, Sedum, 
and Sanicula. 


62 


GARTH AND TILDEN 


/. Re$. Lepid. 


Life Zones: Transition, Canadian, Hudsonian. Plant Communities: 
Juniper Woodland. Host Plants: UMBELLIFERAE: Pteryxia ( = 
Cymoptems) terebrinthina in the Sierra Nevada (Emmel and Emmel, 
1963). 

109 . Papilio rutulus Luc. 

THE WESTERN TIGER SWALLOWTAIL is a species that 
transcends zonal limits in its adherence to the stream-bank associa- 
tion. It occurs at El Portal on the Merced River, thence through the 
Yosemite Valley and its tributary streams well into Canadian Zone. 
One of us (Tilden) has even observed it in Subalpine Forest at Tioga 
Pass. 

Life Zones: Unrestricted. Plant Communities: Willow, Cotton- 
wood, and Aspen groves. Riparian Association. Host Plants: SALI- 
CACEAE: Populus (Cottonwood), Salix (Willow). BETULA- 
CEAE: Alnus (Alder). ROSACEAE: Prunus (Choke-Cherry). 

110. Papilio multicaudatus Kby. 

Syn. Papilio daunus Bdv. 

THE TWO-TAILED SWALLOWTAIL has been known to inhabit 
the Yosemite region since 1930, when B. A. Thaxter and Edna Banta, 
Yosemite Field School members, wrote: "Among the swallowtails found 
here we took Papilio daunus, the largest western species, a bright 
yellow butterfly with black markings and two tails on each hind 
wing.” Their brief report, "Some Butterflies and Moths of the Yose- 
mite Valley Region,” is on file at the Yosemite Museum, but their 
two specimens are without accompanying data. For information on 
exact locality and date we are indebted to Oakley Shields, who twice 
encountered it in August at Jerseydale, where it came to thistles. 

Life Zones: Upper Sonoran, Transition. Plant Communities: All 
at moderate elevations. Host Plants: SALICACEAE: Populus (Cot- 
tonwood), Salix (Willow). LAURACEAE: Umbellularia (Laurel). 
OLEACEAE: Fraxinus (Ash). ROSACEAE: Amelanchier (Service 
Berry), Prunus (= Cerasus) (Choke-Cherry). RUTACEAE: Ptelea 
(Hop Tree). 

Ilia. Papilio eurymedon eurymedon Luc. 

b. Papilio eurymedon albanus F. & F. 

THE PALE SWALLOWTAIL is more frequently encountered on 
dry hillsides than is the nearly related and equally abundant Papilio 
rutulus. While the food plant adheres strictly to the lower zones, the 
strong wings of the butterfly carry it to the tops of the highest peaks. 
A form or possible subspecies occurring at the higher altitudes, smaller 
in size and darker in color, is known as P. e. albanus F. & F. 

Life Zones: Upper Sonoran, Transition, straying higher. Plant 
Communities: Chaparral, Foothill Woodland, and the chaparral-like 
under story of most Coniferous Forests. Host Plants: RHAMNACEAE: 
Rhamnus (= Frangula) calif ornicus (Coffee Berry), R. crocea (Red 
Berry), Ceanothus (Snow Bush). 


2(i):i-^ 6, ip6} 


YOSEMITE BUTTERFLIES 


63 


112. Parnassius clodius haldur Edw. PI. II, fig. e 

THE BALDUR PARNASSIAN flies to the very rim of the preci- 
pitous walls of Yosemite Valley, but does not descend into the valley 
itself. In the Research Reserve it was particularly abundant, flying 
over thickets of Castanea (Chinquapin) and Holodiscus (Ocean Spray), 
and pausing occasionally to sip nectar from yellow Sene do, a composite. 
While the fir belt generally defines its habitat, it occasionally strays 
higher among Lodgepole Pine and Mountain Hemlock. A few indi- 
viduals are without the black-margined secondaries and suggest the 
parent species, P. dodius Men. 

Life Zones: (upper Transition), Canadian, lower Hudsonian 
(Arctic- Alpine ) . Plant Communities: Red Fir Forest; Lodgepole Pine 
Forest; exceptionally, Alpine Fell- Fields. Host Plants: CRASSUL- 
ACEAE: Sedum (Stonecrop) .• ERICACEAE: Vacdnium ( Bilberry 
SAXIFRAGACEAE: Saxifraga (Saxifrage). 

113. Parnassius phoebus hehrii Ed^v. PL III, fig. f 

BEHR’S PARNASSIAN is well established on the eastern slope of 

the Sierra Nevada, as befits its affinities with the Great Basin fauna. The 
arid eastern exposures of many ridges west of the Sierran divide, with 
the persistent association of Juniper and Sedum (Stonecrop), have 
served to lure this butterfly westward through Tioga Pass to Yosemite 
Creek: It also flies in the Potentilla fruticosa zone of the Alpine Rock 
Garden sub-association of the Alpine Fell-Fields. The larvae and pupae 
of P. s. behrii were first described from specimens secured by the 
senior author and Edmund Godwin at Rock Creek Lake. 

Life Zones: Canadian, Hudsonian, Arctic- Alpine. Plant Commun- 
ities: Juniper Woodland, Alpine Fell-Fields. Host Plants CRASSUL- 
ACEAE: Rhodiola rosea (Western Roseroot), Sedum ohtusatum, S. 
stenopetalum (Stonecrop). SAXIFRAGACEAE: Saxifraga (Saxi- 
frage) . 

HESPERIIDAE 

114. Epargyreus darus (Cram.) 

Syn. Epargyreus tityrus ( Fabr. ) 

THE SILVER-SPOTTED SKIPPER is a widely distributed species 
recognizable by its large size and the conspicuous silvery spot on the 
under side of each hind wing. It frequents the flowers of California 
Buckeye, Aesculus calif ornica, a well known tree of the western foot- 
hills, and is favored by plantings of Locust as an ornamental. 

Life Zones: Upper Sonoran. Plant Communities: Foothill Wood- 
land, Yellow Pine Forest. Host Plants: LEGUMINOSAE: Rohinia (In-' 
troduced Black Locust), and other legumes. 

115. Thorybes pylades (Scud.) 

THE NORTHERN DUSKY- WING is the low-elevation species 
of this genus in California. It is larger but has smaller spots on the 
forewing than the following Thorybes mexicana nevada, and the male 
has a costal fold. 


64 


GARTH AND TILDEN 


/. Res. Lepid. 


Life Zones: Upper Sonoran. Plant Communities: Chaparral, Foot- 
hill Woodland, (Yellow Pine Forest). Host Plants: LEGUMINOSAE: 
Trifolium (Clover), and other legumes. 

116. Thorybes mexicana nevadaScwd. 

THE NEVADA DUSKY-WING is the alpine and subalpine 
Thorybes of the Sierra Nevada, flying to 11,000 feet. Smaller and 
with larger spots than the preceding T, pylades, the male has no costal 
fold and the spots are outlined with a dark rim. 

Life Zones: Canadian, Hudsonian, Arctic- Alpine. Plant Commun- 
ities: Alpine Fell-Fields, Subalpine Forest (Glacial Moraine). Host 
Plants: Unknown. LEGUMINOSAE: Amorpha calif ornica (False In- 
digo) for the parent species, according to Comstock and Dammers 
(1933). 

117. Thorybes diversus Bell PL I, fig. a 

BELL’S DUSKY-WING is easily confused with The Northern 

Dusky-Wing, Thorybes pylades, being of similar size, but lacks the 
costal fold in the male. A denizen of grassy openings in the forest belt, 
it shuns the broad, open meadows. 

Life Zones: Transition. Plant Communities: Openings in several 
types of Forest. Host Plants: Not known. 

118. Pyrgus ruralis (Bdv.) 

THE TWO-BANDED SKIPPER prefers openings in the forest 
rather than damp meadows, and is best distinguished by its darker 
color, even checkering, and montane habitat. Flying close to the 
ground, it is inconspicuous and easily overlooked. 

Life Zones: Canadian. Plant Communities: Openings in Red Fir 
and Lodgepole Pine Forest. Host Plants: MALVACEAE: Sidalcea 
( Checkerbloom, Wild Hollyhock). ROSACEAE: Potentilla (Cinque- 
foil) suggested. 

119. Pyrgus communis (Grote) 

THE COMMON CHECKERED SKIPPER is abundant at low 
elevations, often near habitations where introduced mallow grows. It 
may be recognized by the conspicuous flight habit, bluish-gray and 
white checkered pattern, and proximity to human dwellings. 

Life Zones: Upper Sonoran, Transition. Plant Communities: Sev- 
eral. Favors weedy growth around cultivated areas. Host Plants: 
MALVACEAE: especially Malva (Mallow) and Sidalcea (Checker- 
bloom). The larvae draw the leaves together with silken threads. 

120. Heliopetes ericetorum (Bdv.) 

THE LARGE WHITE SKIPPER is a rapid flier, difficult to cap- 
ture, and hence a rarity. The female is more heavily marked than the 
male and resembles the foregoing Pyrgus communis. 

Life Zones: Upper Sonoran (lower Transition). Plant Commun- 
ities: Chaparral, Foothill Woodland, "Mountain Misery” {Chamae- 


2(i):i-96, 1963 


YOSEMITE BUTTERFLIES 


65 


batia folios a) belt of cut-over stands of Yellow Pine Forest. Host 
Plants: AMARANTHACEAE: Amaranthus (Pigweed). MALVA- 
CEAE: Mdvastrum (Apricot Mallow). 

121. Erynnis persius ( Scud . ) 

THE PERSIUS DUSKYWING is a small, dark insect with few 
contrasting markings. Long misidentified in the west, it has been 
shown by Burns (unpublished doctoral dissertation) to be common 
to California at moderate elevations. 

Life Zones: (Upper Sonoran) Transition, Canadian. Plant Com- 
munities: (Foothill Woodland), Yellow Pine and Fir Forests, both 
White and Red. Host Plants: Best regarded as unknown. Early rec- 
ords are untrustworthy. 

122. Erynnis pacuvius lilius ( Dyar ) 

DYAR’S DUSKYWING, as identified by John Burns, is common 
at intermediate elevations. Gin Flat, Crane Flat, and similar locations 
are its usual haunts. One of us (Tilden) found it common on Cow- 
chilla Mountain, outside the park, in June, 1954. 

Life Zones: (upper Transition) Canadian. Plant Communities: 
Chaparral -like understory of montane forests. Host Plants: Unknown. 
RHAMNACEAE: Ceanothus (Snow Bush, Wild Lilac) for Colorado 
pacuvius, as reported by Donald Eff (personal communication). 

123 . Erynnis propertius (Scud. & Burg.) 

THE "PROPERTIUS DUSKWING is the largest and commonest 
of our Duskywings. The front wings have few spots and present a 
slightly "grizzled” appearance, rather than a solid black effect. It 
frequents edges and openings in the forest at moderate elevations. 

Life Zones: (Upper Sonoran) Transition, Canadian. Plant Com- 
munities: Chaparral, Foothill Woodland, Yellow Pine and other Coni- 
ferous Forests. Host Plants: FAGACEAE: Quercus (Oak), especially 
Live Oaks of various species. 

124. Erynnis zarucco funeralis (Scud. & Burg.) 

THE FUNEREAL DUSKYWING is the most easily recognized of 
our Duskywings. The front wing is pointed and the fringes of the 
hind wing are white. There is an indistinct brownish patch on the 
outer third of the forewing that is lacking in the following Erynnis 
tristis; the males also have a tibial tuft. 

Life Zones: (Lower Sonoran) Upper Sonoran. Plant Communi- 
ities: Valley Grasslands, Foothill Woodland (Chaparral). Host Plants: 
LEGUMINOSAE: Hosackia ( = Lotus) scoparia, Medicago (Alfalfa), 
and other legumes. 

125 . Erynnis tristis ( Bdv. ) 

THE SAD DUSKYWING may be expected to occur in the oak 
woodlands and foothills, although it has not yet been found in the 
Yosemite region. Slightly smaller than the preceding Erynnis z. fun- 


66 


GARTH AND TILDEN 


/. Res. Lepid. 


eralis, and with the forewing not noticeably pointed, it lacks the 
brown patch on the forewing and the tibial tuft. 

Life Zones: Upper Sonoran. Plant Communities: Foothill Wood- 
land, Oak Woodland. Host Plants. FAG ACE AE: Quercm (Oak). 

126a. Hesperia harpalus harpalus (Edw.) PL II, fig. g 

b. Hesperia harpalus yosemite Leuss. 

THE HARPALUS SKIPPER is the characteristic Hesperia of the 
eastern declivity of the Sierra Nevada and the adjacent Great Basin 
area surrounding Mono Lake. It is rather pale above, and shows well 
developed silver spots below, especially in the females. THE YOSE- 
MITE SKIPPER is the subspecies that occurs on the western slope 
of the Sierra. It is smaller and more brightly colored than normal 
harpalus and the spotting of the under sides of the hind wings tends 
to be small or to disappear. At high elevations there occurs a "blend- 
zone” form that is intermediate. The literature treats the Great Basin 
form as H. idaho ( Edw. ) , but there seems no reason for retaining this 
name. 

Life Zones: Transition, Canadian. Of the "blend-zone” population: 
Hudsonian ( Arctic- Alpine ) . Plant Communities: Not noticeably se- 
lective. Host Plants: POACEAE: (Grasses). 

127. Hesperia miriamae MacNeill PL III, fig. m 

MIRIAM’S SKIPPER is the most alpine of our skippers, being 

found entirely above tree line on Mono Pass and on the summits of 
the higher peaks, such as Unicorn. It is a small, dark species, with 
large, irregular spots on the under surface of the hind wings. But 
recently described (MacNeill, 1959), it is still a rarity, and will continue 
to be so due to the inaccessibility of its habitat. 

Life Zones: Arctic- Alpine. Plant Communities: Alpine Fell-Fields. 
Host Plants: POACEAE (Grasses) , presumptively. Actually, unknown. 

128. Hesperia nevada (Scud.) 

THE NEVADA SKIPPER is a Great Basin species found on the 
eastern Sierra slope from Mono Lake to elevated areas above Tioga 
Pass. A small species with irregular macular bands on the under side 
of the hind wings, it frequents open rocky areas at high elevations. 

Life Zones: Arid Transition, Canadian, Hudsonian. Plant Com- 
munities: Juniper Woodland, Subalpine Forest, Alpine Fell-Fields. 
Host Plants: POACEAE: Grasses. 

129 . Hesperia juba (Scud.) 

THE YUBA SKIPPER is not limited by elevation, being found 
sparingly across the Sierra crest from the western foothills to the Great 
Basin. A large species of powerful flight, it may be recognized by 
the fuscous border of the front wing, which is sharply limited and does 
not grade into the rufous ground color. The under side of the wings 
is greenish brown to dark brown and the spots are large and usually 
white. 


s(i):i-- 96 > 1963 


YOSEMITE BUTTERFLIES 


67 


Life Zones: Unlimited. Plant Communities: Apparently not re- 
stricted. Ubiquitous. Host Plants: POACEAE: (Grasses), probably. 

130. Ochlodes sylvanoides (Bdy.) 

THE WOODLAND SKIPPER is one of the commonest skippers, 
ranging across the mountains from the western foothills to the Great 
Basin and eastward to Colorado. It is a moderate-sized, orange-brown 
species, with the stigma of the male touching the dusky border of the 
wing. An unwary species, it may be found fairly swarming on roadside 
flowers. The Mono Lake population is slightly paler, but differs from 
Colorado napd Edw., a subspecies with which it has been confused. 

Life Zones: Upper Sonoran, Transition (Canadian). Plant Com- 
munities: Several. Not restricted within its life zones. Host Plants: 
POACEAE : ( Grasses ) . 

131. Ochlodes agricola ( Bdv. ) 

THE FARMER flies from May in the western foothills to July at 
higher elevations, but is absent from the east side of the Sierra Nevada. 
It frequents streamsides, openings in lush woodlands, and overgrown 
roadsides. Smaller and darker than the preceding Ochlodes sylvanoides, 
it shows pale translucent spots in the forewing, at the end of the stigma 
in the male, and near the end of the cell in the female. It is seldom 
found in wet meadows or dry fields. 

Life Zones: Upper Sonoran, Transition. Plant Communities: Open- 
ings in Forests. Riparian and Roadside Associations. Host Plants: 
POACEAE: (Grasses). 

132a. Polites sabuleti sabuleti (Bdv.) 

b. Polites sabuleti tecumseh (Grin.) PL III, fig. d 

THE SANDHILL SKIPPER is found in the Yosemite area, in one 
form or another, entirely across the Sierran crest, from the western 
foothills to the Great Basin. About the size of The Woodland Skipper, 
Polites s. sabuleti has the under surface of the hind wings tan, the spots 
yellowish and extending onto the wing veins, which are also yellow. 
At higher elevations it is replaced by THE TECUMSEH SKIPPER, 
which fairly swarms in the subalpine meadows adjacent to Tioga Pass. 
It is much smaller, darker, brighter, and more greenish than normal 
Polites sabuleti. Its flight season, July to September, is long for an 
alpine species, perhaps indicating two broods. 

Life Zones: Upper Sonoran, Transition for P. sabuleti; Canadian, 
Hudsonian, Arctic- Alpine for P. s. tecumseh. Plant Communities: Not 
restrictive for P. sabuleti; Subalpine Meadow, Alpine Meadow and 
Grassland for P. s. tecumseh. Host Plants: CYPERACEAE: Car ex 
filifolia (Sedge). Also recorded as LEGUMINOSAE: Trifolium 
monanthum (Clover). The latter, if correct, is a departure from the 
food preferences of this group, which are characterized as feeding on 
Monocotyledons. 


68 


GARTH AND TILDEN 


/. Res. Lepid. 


133. Polites sonora {Scud.) 

THE SONORAN SKIPPER is the usual mid-summer skipper of 
intermediate elevations in Yosemite National Park. Resembling the 
Woodland Skipper, Ochlodes sylv amides, in size and appearance, it 
is a darker, more olivaceous rufous, with a thicker, more complex 
stigma. Below, the hind wings are olivaceous tan with a light basal 
spot and a band of light spots across the wing. A somewhat lighter 
phase occurs in the higher portions of the Great Basin, as at Mono 
Lake. 

Life Zones: Arid Transition, Canadian, Hudsonian. Plant Com- 
munities: Meadowlands, Freshwater Marsh, Alkali Sink; openings in 
forests of several kinds. Host Plants: Unknown. 

134.. Amhlyscirtes vialis (Edw.) 

THE ROADSIDE SKIPPER is a widely distributed species, found 
in the cooler parts of the United States and southern Canada from 
California to Maine. In the Yosemite region it is a rarity. A dull dun- 
colored species, of small size, with a few small specks near the tip of 
the forewing, purplish gray shades on the under surface, and checkered 
fringes, it frequents cool, moist spots, such as meadow edges and for- 
est clearings, at elevations of from 4,000 to 7,000 feet. 

Life Zones: Transition, Canadian. Plant Communities: Not known 
for the Yosemite region. Host Plants: POACEAE: (Grasses) in the 
eastern United States; has not been reared in the west. 

LITERATURE CITED 

BADE, WILLIAM F. 1924. The life and letters of John Muir. In two volumes. 

Vol. I: vii 399 pp. Houghton Mifflin, Boston & New York. 

BAUER, DAVID L. 1961. Tribe Melitaeini, Checker Spots, Crescents, and 
Patch Butterflies, pp. 127-146. In: Ehrlich, Paul R., and Anne H. How 
to know the Butterflies. 262 pp. Wm. C. Brown, Dubuque, Iowa. 
BROWN, F. MARTIN. 1954. Colorado Butterflies. Part I. Satyridae. 

Proc. Denver Mus. Nat. Hist., No. 3: 1-32. 

1955. Studies of Nearctic Coenonympha tullia ( Rhopalocera, Satyri- 
dae.) Coenanympha tullia inornata Edw. Bull. Amer. Mus. Nat. Hist., 
105 (4): 363-409. 

CLENCH, HARRY. 1961. Tribe Theclini, Hairstreaks, pp. 177-220. Tribe 
Lycaenini, The Coppers, pp. 220-228. /«:Ehrlich, Paul R., and Anne H. 
How to know the Butterflies. 262 pp. Wm. C. Brown, Dubuque, Iowa. 
COMSTOCK, JOHN A. 1927. Butterflies of California. 334 pp., 63 pis. 
Author, Los Angeles.* 

DAVENPORT, DEMOREST. 1941. The butterflies of the satyrid genus 
Coenonympha. Bull. Mus. Comp. ZooL, Harvard, 87(4): 215-349, 

pis. 1-10. 

DICE, LEE R. 1943. The biotic provinces of North America, vii + 78 pp., 
1 map Univ. Michigan Press, Ann Arbor. 

DOS PASSOS, C. F., and L. P. GREY. 1947. Systematic catalogue of 
Speyeria ( Lepidoptera, Nymphalidae) with designations of types and 
fixations of type localities. Amer. Mus. Novitates, No. 1370: 1-30. 
DOWNEY, JOHN C. 1961. Tribe Plebejini, The Blues, pp. 228-242. In: 
Ehrlich and Ehrlich. How to Know the Butterflies. 262 pp. Wm. C. 
Brown, Dubuque, Iowa. 


196} 


YOSEMITE BUTTERFLIES 


69 


EMMEL, THOMAS C., and JOHN F. EMMEL. 1962. Ecological studies of 
Rhopalocera in a High Sierran community — Donner Pass, California. 
1. Butterfuly associations and distributional factors. /. Lepid. Soc., 16: 
23-44. 

— 1963. Larval food-plant records for six western Papilios. /. Res. 

Lepid., 1(3): 191-193. 

ERIKSEN, C. H. 1962. Further evidence of the distribution of some boreal 
Lepidoptera in the Sierra Nevada. J. Res. Lepid., 1(1): 89-93. ^ 

EHRLICH, PAUL R., and ANNE H. EHRLICH. 1961. How to know the 
butterflies, vi^- 262 pp., 524 figs. Wm. C. Brown, Dubuque, Iowa. 
GARTH, JOHN S. 1935a. Butterflies of the Boundary Hill Research Reserve, 
Yosetmite National Park, California. Bull. Sou. Calif. Acad. Sci., 33: 

131-135. 

— — “ 1935b. Butterflies of Yosemite National Park. Ibid., 34: 37-75. 
GRINNELL, JOSEPH, and HARVEY M. HALL. 1919. Life-zone indi- 
cators in California. Proc. Calif. Acad. Sci., 9(2) : 37-67. 

— , and TRACY T. STORER. 1924. Animal life in the Yosemite. 

xviii +752 pp., 62 pis., 65 figs. Univ. Calif. Press, Berkeley. 

GUNDER, JEAN D. 1929. The genus Euphydryas Scud, of Boreal America 
(Lepidoptera, Nymphalidae) . Pan-Pac. Ent., 6(1) : 1-8, pis. 1-12, maps 2. 
HALL, HARVEY M., and CARLOTTA C. HALL. 1912. A Yosemite flora. 

vii + 282 pp. Paul Elder, San Francisco. 

HEMMING, FRANCIS. 1934. Generic names of Holarctic butterflies. 184 pp. 

British Museum (Natural History), London. 

HIGGINS, L. G, 1953. Butterfly collecting in U.S.A. Entomologist, 86(9) : 
207-210, 

HOVANITZ, WILLIAM 1940. Ecological color variation in a butterfly and 
the problem of "protective coloration.” EcoL, 21 (3): 371-380. 

— — 1943. Hybridization and seasonal segregation in two races of a butter- 
fly occurring together in two localities. Biol. Bull. 85: 44-51. 

— — - — “. 1950. The biology of Colias butterflies. I. The distribution of the 
North American species. Wasmann J. Biol., 8(1): 49-75. 

- — - — 1963. Evidence for the species distinction of Pieris protodice and 
Pieris occidentalis. J. Res. Lepid. 2: in press. 

KLOTS, ALEXANDER B. 1936. The interrelationships of the species of 
the genus Lycaena Fabricius (Lepidoptera, Lycaenidae). Bull. Brooklyn 
Ent. Soc., 31(4): 154-170. 

— 1951. A field guide to the butterflies of North America, east of the 

Great Plains, xvi -f 349 pp., 40 pis., 9 figs. Houghton Mifflin, Boston. 
LINDSEY, ARTHUR W. 1942. A preliminary revision of Hesperia. Den- 
ison Unit’. Bull., Jour. Sci. Labs., 37 : 1-50, pis. 1-6. 

— , E. L. BELL, and R. C. WILLIAMS. 1931. Hesperioidea of North 
America. Ibid., 26: 1-142. 

MATTHES, FRANCOIS E. 1930. Geologic History of the Yosemite Valley. 

Geol. Surv. Professional Paper I6O: 1-137, 52 pis. 

MacNEILL, C. DON. 1959. A new species of Hesperia from California 
(Lepidoptera: Hesperiidae) . Wasmann J. Biol., 17(1): 89-94. 
McDUNNOUGH, J. 1938. Check list of the Lepidoptera of Canada and the 
United States of America. Part 1. Macrolepidoptera. Mem. Sou. Calif. 
Acad. Sci., 1: 1-275- 

MUNZ, PHILIP A., and DAVID D. KECK. 1949. California plant com- 
munities. El Aliso, 2(1): 87-105. 1950. (Same title) — -Supplement. 

Ibid., 2(3) ; 199-202. 

NABOKOV, VLADIMIR. 1949. The Nearctic members of the genus 
Lycaeides Hbl. Bull. Mus. Comp. ZooL, Harvard, 101: 479-541. 

OPLER, PAUL, and JERRY A. POWELL. 1961. Taxonomic and distribu- 
tional studies on the western components of the Apodemia mormo complex 
(Riodinidae) . J. Lepid. Soc., 15(3): 145-171. 


70 


GARTH AND TILDEN 


/. Res. Lepid. 


TILDEN, J. W.. 1959. The butterfly associations of Tioga Pass. Wasmann J. 
Biol., 17(2): 249-271. 

VESTAL, A. G. 1914. Internal relations of terrestrial associations. Amer. 
Nat., 48: 413-445. 

WILLIAMS, C. B. 1930, The migration of butterflies. Oliver and Boyd, 
Edinburgh. 

ZIEGLER, J. BENJAMIN. I960. Preliminary contribution to a redefinition 
of the genera of North American hairstreaks (Lycaenidae) north of 
Mexico. /. Lepid. Soc., 14: 19-23. 


YOSEMITE BUTTERFLY RECORDS 

The documented records on which the textual material rests have been 
segregated from the body of the article (Appendix I) because of its length; 
however these localities are of great importance to specialists interested in the 
specific locations where these species may be found. 

West-slope records precede east-slope records and are separated from 
them by a period. Each locality is followed by all subsequent records from 
that locality, first by the same observer, then by other observers. Localities 
are separated by a semi-colon. Apart from these intended departures an 
exact chronology is maintained. 

Following the abbreviated locality and date (day, MONTH, and year), 
the initials of the responsible observer are given. These are John S. Garth 
(JSG), Edmund D. Godwin (EDG), Allan Oakley Shields (AOS), J. Wilson 
Tilden (JWT), and Keith Trexler (KT). Others who contributed an occas- 
ional record are named with the appropriate citation. 


LOCALITIES AND ABBREVIATIONS 

To conserve space in the body of the list a standard set of abbreviations 
has been adopted, (Appendix II) following the practice established by 
entomological reports of a similar nature (Garth, 1935b) : Localities repre- 
sented extend from Briceburg and Buck Meadows at the western approaches 
to Yosemite National Park to Mono Lake at the eastern Gateway, and from 
Conway Summit on the north to Deadmans Summit on the south along U. S. 
Highway 395. (A few localities, such as Cathay and Oakhurst on the west 
side and Bridgeport and Casa Diablo Hot Springs on the east side, transcend 
even these limits.) Some are in the National Park, some are in the surrounding 
National Forests, while some are outside of either. Localities outside Yosemite 
National Park are identified by National Forest (NF), of if outside the 
National Forest, by California County ( Co. ) . Each locality is characterized 
by Life Zone and Plant Community; the division into west slope and east 
slope suffices to indicate the Biotic Province. Elevations (7214, 10,500) 
are in feet and are taken from U. S. D. I. Geological Survey Topographical 
Maps of Yosemite (1948) or of adjacent Quadrangles. These are accurate 
to the nearest 100-foot contour except in the case of mountain peaks, lake 
levels, or places of habitation, where precise figures (BM) can be given. 


2( i ); i -96, 196} 


YOSEMITE BUTTERFLIES 


71 


APPENDIX I 

SOME PLANTS ON WHICH YOSEMITE BUTTERFLIES FEED AS LARVAE 

o = oviposition observed; 1 - larva taken from; r = reared to adult 
on plant named; ? - doubtful record; * = feeding on more than one 
plant 


AMARANTHAGEAE 

Amaranthus 

ASCLEPIADACEAE 

Asclepias 


Pigweed 


Milkweed 


*Heliopetes ericetorum 


Danaus plexippus o, 1, r 

Danaus strigosus 


BETULACEAE 

Alnus 


BORAGINACEAE 


Amsinckia 

Cryptantha 

Cynoglossum 

Mertensia 


Alder 


*Papilio rutulus 
*Polygonia t. rusticus 


Fiddle-Neck *Vanessa cardui 

Nievitas *Vanessa cardui 

Hound's Tongue *S trymon melinus 

Mertensia Euphydryas c. sierra 


1. r 


1* r 

1, r 


CAPPARIDACEAE 

Cleome 

Isomeris 

CAPRIFOLIACEAE 

Symphoricarpus 
CARYOPHYLLACEAE 
S tellaria 
CHENOPODIACEAE 


Bee Plant *Pieris protodice 

Bladder Pod *Pieris beckeri 


Chickweed 


*Nathalis iole 


o, 1, r 
o, 1, r 


Snow Berry * Euphydryas chalcedona o, 1, r 


Atrlplex 
Chenop odium 


COMPOS ITAE 


Salt Bush *Brephidium exilis 

Lamb's Quarters *Brephidium exilis 


Anaphalis 

Antennaria 

Artemesia 

Aster 


Carduus 


Everlasting 

Everlasting 

Mugwort 

Aster 


Thistle 


^Vanessa virginiensis 1 

*Vanessa virginiensis 1 

*Vanessa virginiensis 
*Melitaea acastus 
? Melltaea d. malcolmi 
Phyciodes campestris 

*Phyciodes mylitta o, 1, r 

* Vanessa cardui o, 1, r 


72 


GARTH AND TILDEN 


/. Res. Lepid. 


Cirsium 

Dyssodia 

Gnaphalium 

Helenium 

Silybum 

CONVOLVULACEAE 

Cuscuta 


CORNACEAE 

Cornus 


CRASSULACEAE 

Rhodiola 

Sedum 


CRUCIFERAE 

Arabia 


Barbarea 

Brassica 


Caulanthus 

Dentaria 

Erysimum 

Lepidium 

Radicula 

Raphanus 

Sisymbrium 


Stanleya 

Streptanthus 

Thlaspi 


Thistle 

(none) 

Cud Weed, 
Everlasting 
Sneeze Weed 
Milk Thistle 


*Phyciodes mylitta 
^ Vanessa cardui 
*Nathalis iole 
^Vanessa virginiensis 

*Nathalis iole 
*Phyciodes mylitta 


o, 1, r 
o, 1, r 

1 


Dodder 


*? Callophrys iroides 


Dogwood 


^ Celastrina argiolus 1, r 


Roseroot *Parnassius 2* behrii 1 

Stonecrop * Parnassius c. baldur 

* P amass jus 2* behrii 1 


Rock Cress 


Winter Cress 
Mustard 


Rock Cabbage 
Milk Maids 
Wall Flower 
Pepper Grass _ 

Water Cress 
Radish 

Hedge Mustard 


Desert Plume, 
Prince's Plume 
Jewel Flower 


Anthocharis lanceolata 
* Anthocharis sara 
^• Euchloe ausonides 
^Euchloe creusa 
*Pieris occidentalis 
* Pieris protodice 
*P_ieris sisymbrii 
*Pieris napi 
* Anthocharis sara 
* Pieris beckeri 
* Pieris napi 
*Pieris protodice 
* Pieris rapae 
*Pieris sisymbrii 
*Pieris napi 
^ •Euchloe ausonides 
^• Pieris occidentalis 
*Pieris protodice 
^Pieris rapae 
*Pieris napi 
*Pieris protodice 
* Anthocharis sara 
*Euchloe ausonides 
*Pieris beckeri 
* ^Pieris occidentalis 
* Pieris protodice 
* Pieris sisymbrii 
*Pleris beckeri 

* Euchloe creusa 
* Pieris sisymbrii 
*Pieris protodice 


o 

o, 1, r 
o, 1, r 


o 


O, 1, r 
o, 1, r 
o, 1, r 
o, 1, r 

o, 1 
o, 1 


o 

o 

o. 


1 


o 


1 . 


r 


Penny Cress 


2(i):i-96, 196} 


YOSEMITE BUTTERFLIES 


73 


ERICACEAE 

Arbutus 

Arctostaphylos 

Rhododendron 

Vaccinlum 

FAGACEAE 

Quercus 


GERANIACEAE 

Erodlum 

GUTTIFERAE 

Hypericum 
LABIATAE 
Hyp t Is 
LAURACEAE 

Umbellularia 

LEGUMINOSAE 

Amorpha 

Astragalus 


Dalea 

Glycyrrhiza 

Hosackia 
( - Lotus ) 


Madrono 

Manzanita 

Azalea 

Bilberry, 

Huckleberry* 


*Callophrys iroides 
*Callophrys iroides 
*Celastrina argiolus 

* Polygonia _f . rusticus 
*Polygonia zephyrus 
*Callophrys iroides 
Colias behrii 
*? Pamassius c. baldur 


1 , 


r 


o, 


1 , 


Oak 


Limenitis b. californica 
*Celastrina argiolus 
Erynnis propertius 
Erynnis tristis 
Habrodais grunus 
? Strymon auretorum 
*? Strymon californica 


Filaree 


*Nathalis iole 


St. John's Wort *Strymon melinus 


Bee Sage 


* Strymon melinus 


Laurel *? Papilio multicaudatus 


False Indigo Colias eurydice 

Thorybes mexicana 
Loco Weed, *Colias eury theme 

Rattle Weed 

*Everes amyntula 
*Everes comyntas 
*Glaucopsyche lygdamus 
* Hemiargu8 isolus 
*Lycaeides melissa 

*Plebe jus acmon 
*Strymon behrii 
*Strymon melinus 
(Includes Smoke *Leptotes marina 
Tree) 

Liquorice * Lycaeides melissa 

*? Celastrina argiolus 
*Colias eury theme 
*? Erynnis z. funeral is 
* Glaucopsyche lygdamus 
*Hemiargus isolus 
* Lycaeides melissa 
* Plebe jus acmon 


o, 1 
o, 1 


o 


o, 1, r 


o 


74 


GARTH AND TILDEN 


/. Res. Lepid. 




*Plebeius saepiolus o 
*Strymon behrii 
*Stirymon melinus 

(2nd brood) 

o, 1, r 

Lathyrus 

Sweet Pea 

*Everes amyntula 
*Glaucopsyche lygdamus 


Lupinus 

Lupine 

*?Celastrina argiolus 
*Glaucopsyche lygdamus 
?Lycaeides a. anna 
*Lycaeides melissa 
Phaedrotes piasus 
Plebeius icarioides 
Strymon fuliginosum 
*Strymon melinus 

o, 1 

o, 1 , r 

Medicago 

Alfalfa 

*Colias eury theme 
*Erynnis z. funeralis 
*Leptotes marina 
*Lycaeides melissa 

0 , 1, r 

Melilotus 

Sweet Clover 

*Colias eury theme 

o 

Rob ini a 

Black Locust 

Epargyreus clarus 


Therraopsis 

False Lupine 

*Colias eurytheme 


Trifolium 

Clover 

Colias philodice hagen 
*Colias eurytheme 
*Everes comyntas 
^Hemiargus isolus 
*Plebeius saepiolus o 
?Polites sabuleti 
Thorybes pylades 

li o, 1, r 
o 

(1st brood) 

Vicia 

Vetch 

*Colias eurytheme 
*Everes amyntula 
*Everes comyntas 
*Glaucopsyche lygdamus 

o, 1 

o 

LORANTHACEAE 

Arceuthobium 

Dwarf Mistle- 
toe 

Callophrys iohnsoni 

Callophrys spinetorum 

1, r 

Phoradendron 

Mistletoe 

Atlides halesus 

1, r 

MALVACEAE 

Lavatera 

Tree Mallow 

*Vanessa carye 


Malva 

Mallow 

*Strymon melinus 
*Vanessa carye 
*PyrKus communis 

r 

o, 1, r 

1, r 

Malvastrum 

Apricot Mallow 

*Heliopetes ericetorum 


Sida 

Alkali Mallow 

*Vanessa carye 


Sidalcea 

Checkerbloom, *Pyrsus coranunis 

Wild Hollyhock 

*?Pyrgus ruralis 
*Vanessa carye 

o, 1, r 

Sphaeralcea 

Bush Mallow, *Vanessa carye 

Apricot Mallow 


ONAGRACEAE 

Gayophytum 


*Lycaena helloides 



2(i):i-^6, 1^6} 


YOSEMITE BUTTERFLIES 


75 


OLEACEAE 

Fraxlnus 
Pit; ACE AE 

Libocedrus 

Pinus 

Pseudotsuga 

PLANTAGINACEAE 

Plantago 

POLYGONACEAE 

Erlogonum 


Oxytheca 

Polygonum 

Rumex 


RHAMNACEAE 

Ceanothus 


Rhamnus 

(= Frangula ) 

ROSACEAE 

Amelanchier 

Cercocarpus 


Ash 


*Papllio multicaudatus 


Incense Cedar 
Pine 

Douglas Fir 


Callophrys nelsoni , probably 
Callophrys eryphon 
^ Neophasia menapia 

*Neophasia menapia 


Plantain 


Euphydryas edltha o, 1, r 

^Precis o. evarete o, 1, r 


Buckwheat 


(none) 
Knot Weed 


Dock, Sorrel 


Apodemia mormo 
Callophrys dumentorum o 

? Callophrys lemberti 
Lycaena gorgon o 

Lycaena heteronea 
? Lycaena nivalis 
Philotes battoides 
Philotes enoptes o 

*Plebeius acmon 
? Plebeius lupini 
*Lycaena helloides 
Philotes speciosa 
*Lycaena helloides 

^Lycaena mariposa 

*S trymon melinus 
*Lycaena editha o 

^Lycaena helloides o 

^Lycaena mariposa 
? Lycaena hypophlaeas 

? Lycaena rubidus 
Lycaena xanthoides o 


Snow Bush 


Cascara, Buck- 
thorn, Coffee 

Berry 


^Callophrys iroides 
*Celastrina argiolus 
Erynnis pgcuvis 
*Papilio eurymedon 
Nymphalis califomica 
S trymon califomica 
S trymon saepium 
*Papillo eurymedon 


1, r 
1, r 


1 

1, r 
1, r 
1, r 


Service Berry *Papillo mul t icaudatus 
Mountain Maho- S trymon adenos,tomatis 1, r 

gany 

S trymon califomica 1, r 

S trymon saepium 


76 


GARTH AND TILDEN 


/. Res. Lepid. 


Potentilla 
(= Horkelia) 

Cinquefoil 

*Lycaena editha 

*?Pyrgus rural is 

Prunus 

(= Cerasus) 

Choke Cherry 

*Limenitis lorquini 
*Papilio multicaudatus 
*Papilio rutulus 

Purshia 

Antelope Bush 

Strymon fuliginosum 

Spiraea 

Spirea 

*Celastrina argiolus 

RUTACEAE 

Ptelea 

SALICACEAE 

Hop Tree 

*Papilio multicaudatus 


1 

1, r 


Populus 


Salix 


Poplar 


Willow 


*Limenitis lorquini 1 

*L imenitis w. nevadae 
*Nyinphalls antiopa o, 1, r 

*Papilio multicaudatus 
*Papilio rutulus o, 1, r 

*Limenltls lorquini 
*Limenitis w. nevadae 
*Nymphalis antiopa o, 1, r 

*Papilio multicaudatus 
*Papilio rutulus 
*? PolyRonia _f. rusticus 

Strymon dry ope 1, r 

S trymon sylvinus 1, r 


SAP INDACE AE 


Aesculus Horse Chestnut *Celastrina argiolus 1, r 

(includes Buckeye) 

SAXIFRAGACEAE 


Glossularia Gooseberry 

Ribes Currant 

SaxigraRa Saxifrage 

SCROPHULARIACEAE 


*Lycaena arota o, 1, r 

*PolyRonia _f . rusticus 
*PolyRonia zephyrus 
*Lycaena arota 
*Pamassius c. baldur 
*Pamassius 2 * behrii 


Castilleja 

Cordylanthus 

Diplacus 


Mimulus 


Penstemon 

Scrophularia 


Paint Brush 

Bird's Beak 
Sticky Monkey 
Flower 

Monkey Flower 

Penstemon 
Figwort, Bee 
Plant 


*Melitaea acastus 
*Melitaea leanira 
*Melitaea leanira 
* Euphydryas chalcedona o, 1, r 

*Euphydryas chalcedona 
^Precis o. evarete 
*Euphydryas chalcedona 
* Euphydryas chalcedona o, 1, r 


2( i ): i -^6, 1963 


YOSEMITE BUTTERFLIES 


77 


ULMACEAE 

Ulmus 

UMBELLIFERAE 

Daucus 

Foenlculum 

Heracleum 

Pteryxia 

URTICACEAE 

Boehmeria 

Parietaria 

Urtica 


VIOLACEAE 

Viola 


Elm 


^Nymphalis antiopa o, 1, r 


Wild Carrot 
Fennel, Anise 
Cow Parsnip 


*Papilio zelicaon 
^Papilio zelicaon 
^ 'Papilio zelicaon 
Papilio indra 


o, 


1, r 
1, r 
1, r 


False Nettle 

Pellitory 

Nettle 


^ 'Vanessa atalanta 
^ Vanessa atalanta 
Nymptialis milberti 
Polygonia satyrus 
^Vanessa atalanta 


1, r 

1, r 


Violet 


Argynnis (all) 
Boloria epithore 


THREE ANOMALOUS SITUATIONS 

While the regularity of zonal progression and the exclusiveness of biotic provinces have 

been emphasized, certain irregularities occur that are of sufficient interest to the lepidopterist 
to warrant considration. The first of these may be found at Tioga Pass, where the loosely 
consolidated moraines provide a situation duplicating the alpine rock garden. Within a few 
feet of these moraines may be seen flying the Arctic-Alpine butterflies, Oeneis chryxus ivallda 
and Euphydryas editha nubigena, species that normally occur above timber line, a thousand 
or more feet higher. The explanation is believed to be found in drainage, both of air that 
funnels through the pass and cools the ground sufficiently to permit the growth of Alpine 

Willow, and of water that enables these slightly elevated rock piles to become snow-free with 
the first rays of summer sun, and to flower a month before the Alpine Fell-Fields above. 

A second paradoxical situation occurs at timberline, as along the Gaylor Lakes trail 
above Tioga Pass, where at an approximate elevation of 10,500 feet may be found Artemesian 
species that normally occur at Mono Lake, 4,000 feet below. Cercyonis oeta, Lycaena rubidus, 
L. heteronea, and Hemiargus isolus are found in a sage-brush environment much like Mono 
Lake, but with Artemesia rotbrockii , rather than A. tridentata, as the sage, and with Eriogonum 
latifolium and Aplopappus sp. as the plant associates. The butterflies are not "fly-ups” from 
below, but appear to be permanently established on these brushy summits. Here the tie to | 
plant community, the Sagebrush Scrub, seems stronger than the restriction to a particular life 
zone or altitudinal belt. Thus one may, after observing "Arctic-Alpine” butterflies at Tioga 
Pass, nominally Hudsonian, ascend a thousand feet and observe "Upper Sonoran” butterflies 
flying at tree-line! 

While normally the Sierran Crest divides the Californian biotic province from the 
Artemesian, a series of lesser crests to the westward support Western Juniper on their east- 

ern slopes, and with it the Juniper-Woodland butterflies, Parnassius phoebus behrii, Euchloe 
ausonides coloradensis, and Hesperia nevada. It is therefore not necessary to travel to the 
eastern slope of the Sierra Nevada to observe Great Basin butterflies; many of these have 

infiltrated the mountain passes and may be found in favored situations as far west as Yosemite 
Creek, well within the central portion of the park. 


78 


GARTH AND TILDEN 


/. Res. Lepid. 


APPENDIX II 
YOSEMITE RECORDS 

SATYRIDAE 

la, Coenonympha tullia californica West. & Hew. El Portal 28- VI- 33 

(JSG); Mather 12- VII- 56 (JSG), 14-V-61 (JWT); Crane Flat 18- VIII- 57 
(JWT); Jerseydale 30-V-59 and 11- VI-61 (JWT), 26- VI-59 (JSG), 30- 
VIII-60 (AOS); Briceburg 21-III-61 (KT); Cliff House 9-VI-61 (JWT). 
b. Coenonympha tullia mono Burdick Mono Lake 3 to 5- VII- 38 and 23- 
VI-62 (JWT), 30-VI-'5Frnd 12- VI- 62 (AOS); W of Lee Vining 26 to 28- 

VI- 61 (AOS, common). 

2. Cercyonis pegala ariane (Bdv. ) Mono Lake 17-VII-57 (JWT), 16 to 20- 

VII- 58 and 13- VII- 60 (AOS); 10 miles E of Mono Lake 24- VII- 59 (AOS). 

3. Cercyonis silvestris (Edw. ) El Portal 28- VI- 33 (JSG), 10- VI- 61 
(JWT); Pate 23-VII-34 (EDG); Benson 24-VII-34 (EDG); Hetch-Hetchy 
13 to 15-VII-56 (JSG); Indian Flat 30-V-59 (JWT, seen), lO-VI-61 
(JWT); Jerseydale 26- VI-59 (JSG), ll-VI-61 (JWT); Cathay ll-VI-61 
(JWT). 

4. Cercyonis oeta( Bdv. ) Tioga Pass 19-VIII-52 (JWT); Crest W of Tioga 
Pass 6-VIII-57, 31- VII to 20- VIII- 58 and 23 to 28- VII- 60 (AOS); Upper 
Gaylor Lakes l-VIII-58 (JSG), 20-VIII-58 (AOS); Pilot Peak 31-VIII- 
58 (JWT). Bodie 18- VII-57 (AOS); June Lake 14 to 22-VIII-57 (JSG); 
Mono Lake 6-VII-58 (AOS), 23- VI-62 (JWT); Warren Creek 2 1 to 28- 
VII-58, ll-VIII-60 and 26-VI-61 (AOS); Agnew Pass and Thousand Is- 
land Lake 5-VIII-59 (AOS). 

5. Oeneis chryxus ivallda (Mead) Dana, W slope 7- VII- 3 1 (JWT); Flor- 
ence 5-VIII-33 (JSG); Lyell 6-VUI-33 (JSG); Dana 8- VIII- 33 (JSG); 

Slide l-VIII-34 (EDG); Pilot Peak 16-VIII-52 and 18- VII-57 (JWT); 

Crest W of Tioga Pass 12 to 15-VII-57 and 9-VII to 19-VIII-58 (AOS); 
Tioga Pass lO-VII-58 (AOS); Helen Lake 18 to 29-VII-58 (AOS); Cocks- 
comb 26-VII-58 (AOS); Dana, N slope 3 1-VII-58 (AOS); Upper Gaylor 
Lake l-VIII-58 (JSG); Vogelsang Pass 3-Vm-58 (JSG); Bert Lake 

9- VIII- 58 (AOS); Unicorn 12- VIII- 58 (AOS). 

DANA IDA E 

6. Danaus (Danaus) plexippus (Linn.) Camp 9, Yosemite Valley, 23-VII- 
33 (JSG); Yosemite Creek 7-VII-56 (JSG); Mather 11 and 12- VII- 56 
(JSG); Crane Flat 19- VII- 57 (JWT); Darrah, Jerseydale, and Indian 
Flat 29 and 30-V-59 (JWT, seen); Jerseydale and Cathay 11- VI-61 
(JWT). Mono Lake 30- VI-41 (JSG, seen), 23-VI-62 (JWT). 

7. Danaus (Tasitia) gilippus strigosus (Bates) Mono Lake 30-VI-30 (JWT, 
seen); l6-Vn-58 (AOSK 

NYMPHALIDAE 

8. Argynnis (Semnopsyche) cybele leto Behr Camp 9, Yosemite Valley, 
23- VII to 2 -VIII- 3 3 (JSG)' 

9. Argynnis (Speyeria) nokomis apacheana Skin. Mono Lake 13- VIII- 50 
and 15- VIII- 51 (JWT), 22 to 26- VIII- 57, 25-VIII-58, and4-VIII-60 
(AOS); Gull Lake 14 to 28-VIII-57 (JSG), 2-IX-58 (JWT). 

10 a. Argynnis (Speyeria) zerene zerene Bdv. Wawona 20- VII- 33 (JSG); 

Jerseydale 21 to 24- VI-56, 19- VI-58, and 9 to 13- VI- 62 (AOS); Mather 
11 to 15-VII-56 (JSG); Hetch-Hetchy Summit 13-VII-56 (JSG); Aspen 


2(i):i~^6, 1963 


YOSEMITE BUTTERFLIES 


79 


Valley 14-VII-56 (JSG); Yosemite Creek Trail 17-VII-56 (JSG); Crane 
Flat I 9 -VIII -57 (JWT). 

b. Argynnis (Speyeria) zerene malcolmi Comst. Agnew Pass 13- VIII- 57 
and 5- VIII - 59 (AOS);“M^^I^Lake 26-VIII-57 (AOS); Lundy 20- VII- 58 
(AOS); Warren Creek 28-Vni-58 (AOS); Casa Diablo 2-IX-58 (JWT). 

11a. Argynnis (Speyeria) callippe inornata Edw. Jerseydale 2 1- VI- 56, 23- 
VI-57, 5- VII- 60, Tl to 23 -VI- 61 and^ to 13- VI- 62 (AOS); El Portal 
30-V-58 (JWT); Oakhurst, Madera County, 24. VI- 62’ (JWT). 
b. Argynnis (Speyeria) callippe nevadensis Edw. Mono Lake 15- VII- 38, 

30 -VI- 50 and 23-VI-62 (JWT), 7-VII-58 (AOS); Above Crowley Lake 

29 - VI-41 (JSG); Bodie 18- VII- 57 (AOS); 5 miles N of Mono Lake 28- VI- 
61 (AOS). 

12. Argynnis (Speyeria) egleis Bdv. Eagle Peak l-VII-33 (JSG); Reserve 
l6^Vn-33 (TsG); Dana 8-Vni-33 (JSG); Pate 23-VII-34 (EDG); Yose- 
mite Creek 9- VII- 56 (JSG); Mather 12- VII- 56 (JSG); Hetch-Hetchy Sum- 
mit 13-VII-56 (JSG); Smoky Jack 4-Vn-57 (JWT); Gin Flat 12 to 19-VII- 

57 (JWT); Lembert Dome 24- VII- 57 (AOS); Crest W of Tioga Pass 6- 
Vni-57, 19-VIII-58, DVni -59 and 14-VIIto 2-VIII-60 (AOS); Crane Flat 

18- VIII-57 (JWT); Pilot Peak 30 to 31-VIII-58 (JWT); Unicorn 27- VII- 
59 (AOS); Glen Aulin 25-VII-60 (AOS). June Lake 14-Vni-57 (JSG); 
Lundy 20-VII-58 (AOS); Warren Creek 21- VII- 58 (AOS). 

13. Argynnis (Speyeria) atlantis irene Bdv. Coldwater 3- VIII- 34 (EDG); 
Tenaya 4- Vni-34 (EDG); Aspen 14- VII- 56 (JSG); 5 miles W of Lower 
Ottoway Lake 19- VIII- 61 (AOS). 

14. Argynnis (Speyeria) hydaspe Bdv. Camp 19, Yosemite Valley 5-Vn- 
33 (JSG); Ledge 9-VII-33 (JSG); Glen Aulin lO-VIII-33 (JSG); Pate 23- 
Vn-34 (EDG); Buck Meadow 4- VII- 54 (JWT); Crane Flat 4- VII- 54 (JWT); 
Jerseydale 28- VI-56, 15 to 16- VI- 58, 18- VI and 4- VH- 60 (AOS), 26- VI- 
59 (JSG); Mather 12-Vn-56 (JSG); Hetch-Hetchy Summit 13 to 17-Vn- 

56 (JSG). 

15. Argynnis (Speyeria) mormonia arge Stkr. Tioga Pass 13-VIII-29 (JSG), 
I4-VII-31, 12 to 13-Vin-50, and“T6^ VIII- 57 (JWT), 4-Vni-57, 19-VIII- 

58 and l-VIII-59 (AOS); Dana 8-Vin-33 (JSG); Coldwater 3-Vm-34 
(EDG); Crane Flat 19-VII to 19-VIII-57 (JWT); Tuolumne Meadows 18- 
Vn to 14-Vni-57 and 3-IX-58 (JWT), 15 to 21-VIII-58 (AOS); Pilot 
Peak 15- VIII- 57 and 3- VIII- 58 (JWT); Dana, W slope l6-Vni-57 and 

30- VIII-58 (JWT); Lyell Fork lO-VIII-58 and 3-VIII-59 (AOS); Gaylor 
Lakes 31-VIIIto 3-IX-58 (JWT); Elizabeth Lake 27-VII-59 (AOS). 

Warren Creek ll-Vni-60 (AOS). 

16 . Boloria (Clossiana) epithore (Edw.) Tioga Pass 7-VII-31 (JWT); Eagle 
Pi"ak I-VIlTFTjSG); LedgT 9-Vn-33 (JSG); Reserve 16-VII-33 and 7 
to lO-VII-56 (JSG); Yosemite Creek 7 to 17-VII-56 (JSG); White Wolf 

10- VII-56 and 24- VI-59 (JSG); Aspen Valley 14-Vn-56 (JSG); Crane 
Flat l9-Vn-57 and lO-VI-61 (JWT), 24- VI-61 (AOS, abundant); Tenaya 

11- VII-58 (AOS); Badger Pass 23- VI-59 (JSG); Glacier Point 23-VI-59t 
(JSG); Tamarack 24- VI- 59 (JSG). 

17a. Euphydryas chalcedona chalcedona (Dbldy. & Hew, ) El Portal 28- VI- 
33 (JSG), 14-V-$1 (JWT) ; Glacier 9-VII-33 (JSG); Jerseydale 16 to 

19- VI-57, 15 to 19-VI.58 and 9 to 13-VI-62 (AOS); Darrah and Jersey- 
dale 30-V-59 (JWT); Indian Flat 30-V-59 and 14-V-61 (JWT). 

b. Euphydryas chalcedona sierra (Wgt. ) Return Creek ll-VII-31 (JWT); 
Eagle Peak l-VII-33 (JSG); Reserve 14-VII-33 (JSG). 


80 


GARTH AND TILDEN 


/. Res. Lepid. 


18a. Euphydryas editha rubicunda Hy. Edw. El Portal 8-V~32 (R. G. Wind), 
14-V-61 (JWT); Jerseydale 15 to 18- VI-58 and 9 to 13- VI- 62 (AOS);in- 
dian Flat 30-V-59, 27-V-60 and 13-V-61 (JWT). 

b. Euphydryas editha nubigena (Behr) Tioga Pass 7-VII-31, 19-VII-52 
and 18-Vn-57 (JWT), 29-VII-56, 8 to 13-VII-57, 9- VII to 24- VIII- 58, 

20 to 23-VII-60 and 25-VI-61 (AOS), l-Vni-58 (JSG); Pilot Peak 18-VII 
to 15-Vm-57 (JWT); Helen Lake 18 to 29-VII-58 (AOS); Gaylor Lakes 
Trail 19-VII-58 (JWT); Dana, W slope 20-Vn-58 (JWT); Rafferty Creek 
3-VIII-58 (JSG); Vogelsang Lake 3-VIII-58 (JSG). 

c. Euphydryas editha monoensis Gund. Mono Lake, near Lee Vining 4- 
VII-28 and 30-VI-50 (JWT); 19-VI-54 (JSG), 30- VI to 16-VII-58, 13-Vn- 

60 and 26 to 27-VI-61 (AOS). 

19. Melitaea (Chlosyne) damoetas malcolmi Comst. Dana, W slope 17- 
VII-3 1 (JWT); Lyell 6- VIII- 33 (JSG); Dana 8- VIII- 3 3 (JSG); Pilot Peak 

16- VIII-52 and 18-Vn-57 (JWT); Upper Gaylor Lake 3 1-VII-57 and 9 
to 19-VII-58 (AOS), l-VIII-58 (JSG); Crest W of Tioga Pass 9-Vnto 

3 1-VII-58, 14-VIIto 2-VIII-60 and 25-VI-61 (AOS); Helen Lake 18-VII-58 
(AOS); Dana, N slope 3 1- VII to 24-Vin-58 and 6 to 8- VIII- 60 (AOS); 

Bert Lake 9- VIII- 58 (AOS). East above Saddlebag Lake 28- VII -58 
(AOS). 

20. Melitaea (Chlosyne) acastus Edw. Mono Lake 5-VII-38 and 30-VI-50 
(JWT), 30-VI-41 (JSG), 30-VI-58 (AOS). 

2 1a. Melitaea (Chlosyne) palla palla Bdv. Eagle Peak 1- VII- 33 (JSGU Ledge, 
Glacier 9-VII-33 (JSG); Reserve l6-Vn-33 (JSG); Yosemite Creek 7 to 

17- VII-56 (JSG); White Wolf lO-VII-56 (JSG); Indian Flat 30-V-59, 27- 

V- 60 and 13-V-61 (JWT); El Portal 14-V-61 (JWT); Jerseydale 21-VI- 

61 (AOS). 

22. Melitaea (Chlosyne) hoffmanni Behr Reserve 14 to 18-VII-33 (JSG); 

Pate 23-VII-34 (EDG); Benson 24-VII-34 (EDG); White Wolf 10-VII- 
56 (JSG). 

23. Melitaea (Chlosyne) leaniraF. & F. Ledge 9-VII-33 (JSG); Tamarack 
Flat 3- VIII- 54 (JWT); Darrah 30-V-57 (JWT); Indian Flat 30-V-59 
and 13-Vto lO-VI-61 (JWT); El Portal 14-V-61 (JWT); Jerseydale 
22-VI-61 (AOS). 

24a. Phyciodes (Phyciodes) campestris campestris (Behr) Museum 5-Vn- 
33 (JSG); Mather 12- VII- 56 (JSG), 10- VI-61 and 30-VI-62 (JWT); 
Hetch-Hetchy 13-VII-56 (JSG); Jerseydale 23-VI-61 (AOS). 

b. Phyciodes (Phyciodes) campestris montana (Behr) Reserve l6-Vn-33 
(JSG); Dana 8-Vni-33 (JSG); Coldwater 3- VIII- 34 (EDG); Tamarack 
Flat 3-VII-54 (JWT); Aspen Valley 14-Vn-56 (JSG); below Helen Lake 
19-VII-60 (AOS). Thousand Island Lake 4-VIII-59 (AOS). 

c. Phyciodes (Phyciodes) campestris, intermediate form Mono Lake 30- 

VI- 41 and 19-VI-54 (JSG), 17- VII- 57 and 23-VI-62 (JWT), 30-VI to 20- 

VII- 58, 13-VII-60, 26 to 27- VI-61 and 12-VI-62 (AOS); Huntoon Public 
Camp, Bridgeport 19- VI- 54 (JSG); June Lake 14 to 29- VIII- 57 (JSG); 
Warren Creek 21 and 28- VII- 58 (AOS); W of Lee Vining 26 to 27- VI- 61 
(AOS). 

25. Phyciodes (Phyciodes) mylitta (Edw. ) Museum 5-VII-33 (JSG); Benson 
24-VII-34 (EDG); Yosemite Creek 9-VII-56 (JSG); Aspen Valley 14- VII- 
56 (JSG); Crane Flat 18-Vm-57 (JWT), 24- VI-59 (JSG); Tioga Pass 1- 
IX-57 (AOS); Tenaya ll-Vn-58 (AOS); Glacier Point 23-VI-59 (JSG); 
Jerseydale 26-VI-59 (JSG), 30- VIII- 60 (AOS); El Portal 2 l-IH- 6 1 (KT), 


2( i ): i . s 6, 1963 


YOSEMITE BUTTERFLIES 


81 


14-V-61 (JWT);Yosemite Valley 19”IV~6l (KT); Indian Flat 13-V-61 
(JWT); Mather 10- VI-61 (JWT); Warren Creek ll-VIII-60 (AOS). 

26. Polygonia satyrus (Edw. ) Museum 5-VII-33 (JSG). Mono Lake 20-VI 
t^25-VIII-58T^27-VI-6l (AOS). 

27. Polygonia faunus rusticus (Edw.) "Yosemite" VI-26 (E, O. Essig). 
Specimen in Yosemite Museum in 1933. 

28. Polygonia zephyrus (Edw.) Reserve 14-Vn-33 (JSG); Yosemite 
^eek 9-'viLT6 (JSG); Aspen Valley 14-Vn-56 (JSG); Jerseydale 17- 
VII-56 and 16-VI-58 (AOS); Dana, W slope 20-VII-58 (JWT); Tioga 
Pass 20-VIII-58 (AOS); Upper Gaylor Lake 20-VIII-58 (AOS); Dana, N 
slope 31-VII to 24-VIII-58 and 8-Vin-60 (AOS); Badger Pass 23-VI-59 
(JSG); Tuolumne Grove 24- VI-59 (JSG). Mono Lake 30-. VI-41 (JSG). 

29 . Nymphalis californica (Bdv. ) Yosemite Valley 24- VI to 5-VII-33 
(JSG); Ledge 9-VII-33 (JSG); Darrah, Jerseydale, and Indian Flat 30- 

V- 59 (JWT); Bridal Veil Creek 23- VI-59 (JSG); White Wolf 24- VI-59 
(JSG); Cliff House 9- VI- 61 (JWT); Crane Flat, Gin Flat, and Mather 
lO-VI-61 (JWT) (outbreak numbers); El Portal lO-VI-61 (JWT); Jer- 
seydale and Cathay 11- VI-61 (JWT). Donohue Pass 4-VIII-59 (AOS). 

30. Nymphalis milberti furcil-lata (Say) Lyell 6- VIII- 33 (JSG); Tioga 
Pass 8-VIIt^-Vin-57, 1- VIII- 58, and 25- VI- 61 (AOS); Crest W of 
Tioga Pass 1 to 19-VIII-58 (AOS); Lyell Base Camp 9-VIII-58 (AOS); 
Dana, N slope 24-VIII-58 (AOS). Mono Lake 30-VI-41 (JSG), 26-VI- 
61 (AOS); Warren Creek 29- VIII- 57 (AOS); Agnew Meadows 9- VIII- 
58 (AOS). 

31. Nymphalis antiopa (Linn.) Research Reserve 8-'VdI-56 (JSG, seen). 
Mono Lake 30- VI-41 (JSG, seen). 

32. Vanessa atalanta (Linn.) "Yosemite" VI-26 (E. O. Essig). 

33. Vanessa virginiensis (Dru. ) Research Reserve 14 to 18-VII-33 and 
T0^^Yn^6 (JSG); Yosemite Creek Trail 9-VII-56 (JSG); Mather 10- 

VI- 61 (JWT). 

34. Vanessa cardui (Linn.) Research Reserve 14 to 18-VII-33 (JSG); 

YosT mit e“cT^ Trail 9- VII- 56 (JSG). 

35. Vanessa carye (Hbn. ) Research Reserve 14 to 18-VII-33 (JSG); 

Hatch- Hetchy 12- VII- 56 (JSG). Huntoon Public Camp, Bridgeport 19- 
VI- 54 (JSG). 

Precis orithya evarete (Cram.) Ledge Trail 9-VII-33 (JSG); Research 
Reserve 14 to 19-VII-33 and 8 to lO-VII-56 (JSG); Yosemite Creek 
Trail 9“ VII- 56 (JSG); Mather Hand 12-VII-56 (JSG), lO-VI-61 (JWT); 
Jerseydale 17~VI-60 (AOS); El Portal 4-IV- 61 (KT); Indian Flat 13- V- 
61 (JWT); Cathay 11- VI- 61 (JWT). Mono Lake 13- VII- 60 (AOS). 

37. Limenitis (Limenitis) weidemeyerii nevadae (B. Er B.) Mono Lake 30- 
VI-41 (jSG)7“30-VI-567jWTT7 30-VI to 20TviI-58 and 26- VI- 61 (AOS). 
Limenitis (L. ) weidemeyerii nevadae X Limenitis (L.) lorqmni (Hy- 
brid) Mono“Lak7l7^VIII- 5 1 (JWT), 19- VI- 54 (JSgJ, 7 to 20- VII- 58, 
25-VII-59 and 27-VI-61 (AOS). 

38. Limenitis (Limenitis) lorquini Bdv. Yosemite Valley VII- 33 (JSG); 

Pate ValIiy'23-VII-34 "(EDCFAspen Valley 14- VII-56 (JSG); Jersey- 
dale 16-VI-58 (AOS); Darrah and Jerseydale 30-V-59 (JWT, seen); 
Indian Flat 30-V-59 (seen) and 13-V-61 (JWT); Cliff House 9-VI-61 
(JWT); Mather lO-VI-61 (JWT). Mono Lake 30-VI-58 (AOS). 

39 . Limenitis (Adelpha) bredowii californica (Butl.) Museum 5-Vn-33 
(JSG); Camp 9, Meadow 23-VII-33 (JSG); Mather 12 to 15- VII- 56 (JSG). 


82 


GARTH AND TILDEN 


/. Res. Lepid. 


10- VI-61 (JWT); Hetch-Hetchy 13 to 15-VII-56 (JSG); Aspen Valley 14- 
VII-.56 (JSG); Crane Flat 24- VI-59 (JSG); Jerseydale 26-VI-59 (JSG), 

11- VI-61 (JWT), 9 to 13-VI-62 (AOS); Darrah, Jerseydale, and Indian 
Flat 30-V-59 (JWT); El Portal 14-V-61 (JWT); Cliff House 9-VI-61 
(JWT). 


RIODINIDAE 

40a. Apodemia mormo mormo F. & F. Little Yosemite 3-VIII-33 (JSG). 
Mono Lake 22 to 26- VIII- 57 and 20- VII- 58 (AOS); June Lake 22-Vm- 
57 (JSG); Conway Pass, N of Mono Lake 21-VIII-60 (JWT). 
b. Apodemia mormo tuolumnensis Opler & Powell Pate Valley 23-VII-34 
(EDG, as A. m. virgulti). 

LYCAENIDAE 

41. Habrodais grunus (Bdv. ) Columbia Point 14-VII-33 (JSG); Pohono 
Trail 23- VII- 33 (JSG); Benson 24- VII- 34 (EDG); Hetch-Hetchy 13- VII- 

56 (JSG); Jerseydale 21 to 23-VI-61 (AOS). 

42. Atlides halesus estesi Clench Museum 5-Vni-34 (collector unknown); 
Yosemite Creek Trail 9- VII- 56 (JSG). 

43. Strymon (Strymon) melinus pudica (Hy. Edw. ) Museum 8-VII-33 
(JSG); Mather 12- VII- 56 (JSG); Darrah 29-V-59 (JWT). 

44. Strymon (Satyrium) fuliginosum semiluna (Klots) Mono Lake 30- VI- 50 

(JWT), 3 O-VI-58 ^ A'osyi 

45. Strymon (Satyrium) behrii (Edw.) Mono Lake 30-VI-50 and 23-VI-62 
(JWT), 30-VI to 20- VII-58 and 27- VI- 61 (AOS); W of Lee Vining 18- 
Vn-57 and 24- VI-58 (AOS); Bodie 18-VII-57 (AOS); Mono Craters 
23- VII- 57 (AOS); Warren Creek 26- VI-61 (AOS). 5 miles N of Mono 
Lake 28- VI- 61 (AOS). 

46. Strympn (Satyrium) auretorum (Bdv.) El Portal 28- VI-33 (JSG), 10- 
VI- 61 (JWT); Briceburg and Mariposa 26- VI- 54 (JWT); Hetch-Hetchy 
Road 13 to 15-VII-56 (JSG); Jerseydale 21 and 22- VI-58 (AOS); Darrah 

29- V-59 (JWT); Cathay ll-VI-61 (JWT). 

47. Strymon (Satyritim) saepium (Bdv.) El Portal 28- VI-33 (JSG); Pate 
23-VII-34 (EDG); Hetch-Hetchy Road 13-VII-56 (JSG); Gin Flat 17-Vni- 

57 (JWT); Jerseydale 21 and 22- VI- 58 (AOS); Darrah and Indian Flat 

30- V-59 (JWT); Cathay 28-V-60 and ll-VI-61 (JWT); Cliff House 9- 
VI- 61 (JWT). Warren Creek 28- VII and 28- VIII- 58 (AOS); Gull Lake 
2-IX-6I (JWT). 

48. Strymon (Satyrium) adenostomatis (Hy. Edw.) Mariposa 26-VI-54 
(JWT); Jerseydale 21 and 22-VI-58 (AOS); Darrah 29-V-59 (JWT). 

49. Strymon (Satyrium) sylvinus (Bdv.) Museum 8-Vn-33 (JSG); Bear 
Creek LodJiT^^I- 5T(TWT); Hetch-Hetchy Road 13-VII-56 (JSG); 
Jerseydale 2 1- VI- 6 1 <(AOS). Mono Lake 5-VII-38 (JWT). 

Strymon ( Satyrium) californica (Edw.) Old Village 12-VI-32 (JWT); 
Museum 5-VII-33 (JSG); Pate 23- VII- 34 (EDG); Big Oak Flat Road 
2-VII-54 (JWT); Mirror Lake 2- VII- 54 (JWT); Tamarack Flat 4-Vn- 
54 (JWT); Hetch-Hetchy Road 13 to 15- VII- 56 (JSG); Mather 15- VH- 
56 (JSG); Jerseydale 15 to 22-VI-58, 17- VI- 60 and 2 1 to 23- VI- 6 1 

(AOS); Darrah 30-V-59 (JWT); El Portal 14- V and 10- VI- 6 1 (JWT); 
Cathay ll-VI-61 (JWT). Mono Lake 5- VII- 38, 30-VI-50, 17-Vn-57, 
and23-VI-62 (JWT), 30- VI-58 and 27- VI-6 1 (AOS); Lee Vining 24- 
VI-58 (AOS); Warren Creek 2 1- VII-58 (AOS); 5 miles N of Mono 


2(i):i-s 6, t$6} 


YOSEMITE BUTTERFLIES 


83 


Lake 28- VI- 61 (A”OS). 

Strymon ( Satyrium) dryope (Edw. ) Mono Lake 20- VII- 58 and 4- VIII- 60 
(AOS); Bridgeport 15- VIII- 52 (JWT). 

52. Callophrys (Callophrys) dumetorum (Bdv. ) Briceburg 21-III- 6 1 (KT); 

El Portal l6TlvT6l"~(JWTTr“ Mather 14-V-61 (JWT). 

53. Callophrys (Callophrys) lemberti Tilden Reserve 16-VII-33 (JSG); 
Pil^ Peak I'o'-VII and l6^Wl-5’2 (JWT); White Wolf lO-VII-56 (JSG); 
Crest W of Tioga Pass 9-VII-58 and 25-VI-61 (AOS), 20-VII-58 
(JWT); Tioga Pass lO-VII-58 (AOS). Warren Creek 12-VI-62 (AOS). 

54. Callophrys (Mitoura) spinetorum (Hew.) Tamarack Flat 4- VII- 54 
(JWT); JersT^^daTI” I?- VI- 57Ti7d 27-VIIto 3- VIII- 62 (AOS); Tioga 
Road 24- VI- 59 (JSG). 

55. Callophrys (Mitoura) johnsoni (Skin.) Jerseydale 27 and 28-VII-56, 
^VIII- 56 anT^T^Vn-I^lAOS) . 

56. Callophrys (Mitoura) nelsoni (Bdv.) Musetim 5-VII-33 (JSG); Reserve 
Ti toT8-'^L3T7jSG)rijetch-Hetchy 13-VII-56 (JSG); Jerseydale 30- 

V- 59, 27-V-60, and ll-VI-61 (JWT), 26- VI-59 (JSG), 9 to 13-VI- 

62 (AOS); Crane Flat 24- VI-59 (JSG); Highway 120 and Hetch-Hetchy 
Road 14- V- 61 (JWT); Cliff House 9-VI-61 (JWT); Mather 10- VI- 61 
(JWT). 

57. Callophrys (Incisalia) doudoroffi windi (Clench) Indian Flat 16-IV-61 

__ _ 

58. Callophrys (Incisalia) iroides (Bdv.) Ledge 9-VII-33 (JSG); Crane 
Flat 3- VII- 54 (JWT); “jerseydale 19- VI-58 and 21 to 23- VI-61 (AOS), 
29-V-59 (JWT); Darrah 29-V-59 (JWT); Glacier Point 23-VI-59 

'(JSG); El Portal 4-IV-61 (KT); Mather lO-VI-61 (JWT). 

59. Callophrys(Incisalia)eryphon (Bdv.) Reserve 9 to 14-VII-33 (JSG); 
TamaraclTFlat 4- VII- 54 (JWT); Smoky Jack 4- VII- 54 (JWT); 

Mather 12-VII-56 (JSG), lO-VI-61 (JWT); Jerseydale 16-VI-58 
(AOS), 29-V-59 (JWT); Tenaya Canyon 11- VII- 58 (AOS). Mono 
Lake 27- VI- 61 (AOS). 

60a. Lycaena (Tharsalea) arota arota (Bdv.) Museum 8-VII-33 (JSG); 

^te 23-VJl34~l¥DG)r^e77i7dale 29-VI to 27-VII-56, l-VIII-58, 
24-Vin-60, 23-VI-61 and 28-Vn-62 (AOS), 
b. Lycaena (Tharsalea) arota virginiensis (Edw.) Mono Lake 17-VII-57 
TjWtTT” 2 6‘^^IL 5 7T rt^O-VII-58 and 13- VII- 60 (AOS). 

61. Lycaena (Lycaena) gorgon (Bdv.) Darrah 29-V-59 (JWT); Indian 
^at 30 -vT 59~^ ll-V to lO-VI-61 (JWT); El Portal 14-V-61 (JWT). 

62. Lycaena (Lycaena) heteronea Bdv. Lyell Base Camp 26- VII- 57 (AOS); 
Tioga Pas”i~4rvni-“5T"lAOS); Merced Lake Trail 4- VIII- 58 (AOS); 
Gaylor Lakes Trail 3-IX- 58 (JWT); Dana, N slope 8-Vni-60 (AOS). 
Warren Creek 7- VIII-57 and 2 1- VII to 28-Vm-58 (AOS); Lundy 20- 
VII- 58 (AOS); Mono Lake 13- VII- 60 (AOS). Thousand Island Lake 
4-Vm-59 (AOS), 

63. Lycaena (Lycaena) xanthoides (Bdv.) Mather 11 and 12-VII-56 (JSG), 

(JWT);” HSt^^Hitchy 13- VII- 56 (JSG); Aspen Valley 14- 
VII- 56 (JSG). 

64. Lycaena (Lycaena) rubidus (Behr) Mono Lake 27-VII-37 (JWT), 30- 

VI- 41 (JSG), 30- VI to 7^11-58 and 27- VI-61 (AOS); Gaylor Lakes 
Trail 16-VIII-52 (JWT); Warren Creek 7 to 29-Vin-57, 21-VIIto 
28-VIII-58 and ll-FIII-60 (AOS); Agnew Pass 5-Vm-59 (AOS); W 
of Lee Vining 2 8- VI- 61 (AOS). 


84 


GARTH AND TILDEN 


/. Res. Lepid. 


65. Lycaena (Lycaena) editha (Mead) Kuna 7- VIII- 33 (JSG); Dana 8- VIII- 

33 (JSG): Kerrick 27- VII- 34 (EDG); Snow Lake 29- VII- 34 (EDG); 
Tamarack Flat 4- VII- 54 (JWT); Tioga Pas s 16 to 17- VII- 57 (JWT), 

14- VII to 24- VIII- 58 aTid 14- VIII- 60 (AOS); Gin Flat and Crane Flat 
19-VII-57 (JWT); Rafferty Creek 21-VII-57 (AOS); Tioga Meadow 

15- VIII-57 (JWT); Pilot Peak 15- VIII- 57 and 3 1 - VIII- 58 (JWT); 

Dana, W slope 17- VIII- 57 and 20- VIII- 58 (JWT); Tuolumne Meadows 
15 to 21- VIII-58 (AOS), 3-IX-58 (JWT); Gaylor Lakes Trail 30-VIII 
to 3-IX-58 (JWT); Lyell Base Camp 3-VIII-59 (AOS); below Agnew 
Pass 5-VIII-59 (AOS). Mono Lake 30- VI-41 (JSG); Bodie 18-VII-57 
(AOS); Warren Creek 21-VII-58 (AOS). 

66. Lycaena (Lycaena) mariposa Reak. Kuna 7-VIII-33 (JSG); Slide 29- 

VII- 34 (EDG); Dana, W slope 13-VIII-50, 15-VIII-51, 16-VIII-57 

and 30- VIII-58 (JWT); Lyell Base Camp Trail 26-VII-57 and lO-VHI- 
58 (AOS); Tioga Pass 4-VIII-57, 20- VIII- 58 and 1- VIII- 59 (AOS); 

Pilot Peak 15-VIII-57 (JWT); Elizabeth Lake 27- VII to 22- VIII -58 
and 27- VII - 59 (AOS); Merced Lake Trail 4- VIII- 58 (AOS). Agnew 
Meadow 13- VIII- 57 (AOS). 

67. Lycaena (Lycaena) nivalis (Bdv. ) Tioga Pass 14-VII-31 (JWT), 20- 

VIII- 58 (AOS); Glacier 9- VII- 33 (JSG); Reserve 14-VII-33 (JSG); 
Kuna 7- VIII- 33 (JSG); Benson 24-Vn-34 (EDG); Kerrick 27- VII- 

34 (EDG); Gaylor Lakes Trail 15-VIII-51, 6- VIII- 53 and 19- VII to 3- 

IX- 58 (JWT); Smoky Jack 3- VII- 54 (JWT); Yosemite Creek Trail 

9- VII- 56 (JSG); Dog Lake Trail 19- VII- 56 (JSG); Rafferty Creek 
Trail 21- VII- 57 and 24- VII- 58 (AOS); 3-VIII-58 (JSG); Tenaya Can- 
yon ll-VII-58 (AOS); Crest W of Tioga Pass 4 to 18- VIII- 57, 20- 

VIII- 58 and 20 to 23- VII- 60 (AOS); Lyell Fork Meadows 10- VIII- 58 
and 3- VIII - 59 (AOS); Dana, N slope 24- VIII- 58 (AOS). Warren 
Creek 21-VII-58 (AOS). 

68. Lycaena (Lycaena) helloides (Bdv.) El Portal 28- VI-33 (JSG); Mather 
12- VII- 56 (JSG), Mono Lake 5-VII-38 (JWT). 

69 . Lycaena (Lycaena) phlaeas hypophlaeas (Bdv.) Lyell 6-Vin-33 (JSG); 
Dana, N slope 6 to 13- VIII- 60 (AOS). 

70. Lycaena (Lycaena) cupreus (Edw. ) Tioga Pass 7-VII-31, 13-Vni-50, 

15- VIII- 51, 16- VII to 16- VIII- 52, 6-VIII-53, and 18- VII to 16-VIII-57 

(JWT), 7 to 15-Vn-57, 9-VII to 24- VIII-58, 15-VII-60 and 25-VI-61 
(AOS); Kuna 7-VIII-33 (JSG); Dana 8-Vin-33 (JSG); Snow Lake 29- 
VII-34 (EDG); Dog Lake Trail 19-VII-56 (JSG); Pilot Peak 18- VU 
to 15-VIII-57 (JWT); Cockscomb Peak 26- VI-58 (AOS); Tuolumne 
Meadows 4-VII-58 (AOS); Crest W of Tioga Pass 9-VII to 1- VIII- 58 
(AOS); Helen Lake 18 to 29- VII- 58 (AOS); Elizabeth Lake 27-VII- 

58 (AOS); Upper Gaylor Lake 1- VIII- 58 (JSG); Rafferty Creek Trail 
3- VIII- 58 (AOS). 

71. Leptotes marina (Reak.) Mono Lake 30- VI-41 (JSG), 30- VI-50 

(JWT, seen) and 23-VI-62 (JWT). 

72. Brephidium exilis (Bdv.) E above Saddlebag Lake 28-VII-58 (AOS); 
Mono Lake 23-VI-62 (JWT). 

73. Hemiargus (Echinargus) isolus (Reak.) Tioga Pass 9-Vnto 7-Vin-58 
(AOS); Crest W of Tioga Pass 9- VII- 58 (AOS); Upper Gaylor Lake 
14-Vn-58 (AOS); Rafferty Creek Trail 3- VIII- 58 (JSG, AOS); Vo- 
gelsang Lake 3-VIII-58 (JSG, AOS); Gaylor Lakes Trail 31-VIIIto 
3-IX-58 (JWT); Tuolumne Meadows 3-IX-58 (JWT). Mono Lake 20- 
VU-58 (AOS); Lee Vining 24-VII-58 (AOS); Warren Creek 28-VII- 


2( i ): i -96, 1963 


YOSEMITE BUTTERFLIES 


85 


58 (AOS). 

74. Lycaeides argyrognomon anna (Edw. ) Conness Creek ll-VII-31 

■(JWT); Dana 8- VIII- 33 (JSG);" Kerrick 27- VIE 34 (EDG); Coldwater 
3- VIII- 34 (EDG); Tenaya 4- VIII- 34 (EDG); Crane Flat 2- VII- 54 
and 19-Vn-57 (JWT); Hetch-Hetchy Summit 13-VII-56 (JSG); Aspen 
Valley 14-VII-56 (JSG); Gin Flat 19-VII-57 (JWT); Tuolumne Mea- 
dows 14-VIII-57 (JWT); Badger Pass 23-VI-59 (JSG). 

75. Lycaeides melissa inyoensis Nab. Reserve 16-VII-33 (JSG); Ben- 
son 24-Vli3riEbG); Helen Lake 18- VII- 58 (AOS); Cockscomb 
Peak 26-VII-58 (AOS); Upper Lyell Base Camp lO-VIII-58 and 3- 
VIII - 59 (AOS). Mono Lake 20- VII- 58 (AOS), 23- VI- 62 (JWT); E 
above Saddlebag Lake 28-VIL58 (AOS); Warren Creek ll-Vni-60 
and 26- VI-61 (AOS); Conway Sximmit 23-VI-62 (JWT). 

76 . Plebejus (Agriades) glandon podarce (F. & F. ) Tioga Pass 7-VII-31, 
TTand 14 ->111-50, 15 - VIII- 517 16 - VIH- 52, 6 and 7- VIII- 53, and 16- 
VIII-57 (JWT), 1 to 24-Vm-58 and 23- VII-60 (AOS); Eagle Peak 1- 
Vn-33 (JSG); Reserve 18-VII-33 and lO-VII-56 (JSG); Kuna 7-VIII- 

33 (JSG); Kerrick 27-VII-34 (EDG); Tuolumne Meadows 16-VII-57 
(JWT); Dana, W slope 17-VIII-57 and30-Vni-58 (JWT); Elizabeth 
Lake 27-VII-58 (AOS); Helen Lake 29-VII-58 (AOS); Upper Gaylor 
Lake l-VIII-58 (JSG); Bert Lake 9-VIII-58 (AOS); Unicorn Peak 
23-Vni-58 (AOS); Bridal Veil 23- VI- 59 (JSG); White Wolf 24- VI- 

59 (JSG); Dana, N slope 8 -VIII- 6 O (AOS). 

77. Plebejus (Plebejus) saepiolus (Bdv.) Glacier 9- VII- 33 (JSG); Re- 

7^77^~T7- Vn-33 (JSG); Florence 5-Vin-33 (JSG); Pate 23-Vn-34 
(EDG); Tioga Pass 15-VIII-51, 6- VIII- 53 and 16 to 18- VIE- 57 (JWT), 

9- VII to 24- VIII- 58 and 13 to 15- VII- 60 (AOS); Crane Flat 3- VII- 54 

and 19 - VII- 57 (JWT); Hetch-Hetchy Summit 13-VII-56 (JSG); Aspen 
Valley 14- VII- 56 (JSG); Tuolumne Meadows 18- VII- 57 (JWT); Gin 
Flat I 9 -VII -57 (JWT); Dana, W slope 17- VIII- 57 and 20- VII to 30- 
Vin-58 (JWT); Crest W of Tioga Pass 9~Vn to 20-Vni-58 (AOS); 
Helen Lake 18- VII- 58 and 19- VII- 60 (AOS); Rafferty Creek Trail 
3-Vin-58 (JSG, AOS); Gaylor Lakes Trail 3 1- VHI to 3-IX- 58 (JWT); 
Bridal Veil and Badger Pass 23- VI- 59 (JSG); White Wolf, Tioga Road, 
and Crane Flat 24- VI-59 (JSG). Mono Lake 5-VII-38, 30- VI- 50 and 

14-Vni-57 (JWT), 30-VI-41 (JSG), 13-VII-60 (AOS). 

78a. Plebejus (Icaricia) icarioides icarioides (Bdv.) Glacier 9-VII-33 

TjSG)r^si"en Valle7~T4- VII- 56 (JSGF” below Vogelsang Pass 3-Vni- 
58 (AOS); Badger Pass 23- VI- 59 (JSG); Crane Flat 24- VI- 59 (JSG); 
Jerseydale 29 to 30- V- 59 and 1 1- VI- 6 1 (JWT), 26- VI- 59 (JSG), 9 
to 13-VI-62 (AOS); Indian Flat 13- V-6 1 to 10- VI-6 1 (JWT); Mather 

10- VI-61 and 30-VI-62 (JWT). 

b. Plebejus (Icaricia) icarioides hellos Edw, Mono Lake d-VII-SS and 
TJTvLhl (JWT), 37^-41 and 19-VI-54 (JSG), 7 to 20- VII- 58 and 
26 to 27- VI-61 (AOS); Conway Pass, ‘ N of Mono Lake 30- VI-50 and 
23-VI-52 (JWT); Huntoon Public Camp, Bridgeport 19- VI-54 (JSG); 
Bodie 28- VII- 57 (AOS); Agnew Pass and Donohue Pass 4-Vni-59 
(AOS); Warren Creek ll-VIII-bO (AOS); W of Lee Vining 26- VI- 61 
(AOS); 5 mi N of Mono Lake 28- VI-61 (AOS). 

Plebejus ( Icaricia ) shasta comstocki Fox Reserve l6-Vn-33 (JSG); 
Kuna 7-Vn-33 (JSG); Ke rTI^ 2 7^ VII- 3 4 (EDG); Slide Creek 29- VE- 

34 (EDG); Tioga Pass 13-VIII-50, 15-VEI-52, 5 to 6-VEI-53, and 

16-Vin-57 (JWT), 15-Vn-57, 14-Vn to 20-Vm-58 and 15 to 20-VE- 


86 


GARTH AND TILDEN 


/. Re$. Lepid. 


60 (AOS); Pilot Peak 15- VIT 57 and 19- VH to 3 H VIII- 58 fJWT); Gin 
Flat 17-Vni-57 (JWT); Dana, W slope 17-VIII-57 and 30-Vni-58 
(JWT); Upper Gaylor Lake 14 to 19- VII- 58 (AOS); Helen Lake 18- VH- 
58 (AOS); Upper Lyell Base Camp 9-VnL58 (AOS); Crest W of 
Tioga Pass 19 to 20-Vni-58 (AOS); Dana, N slope 24-Vin-58 and 8- 
VIII-60 (AOS); Gaylor Lakes Trail 3-IX-58 (JWT). Bodie 18-VIL 

57 (AOS); Lundy ZO-VII-58 (AOS); Donohue Pass 4 to 6~ VIII- 59 
(AOS); Warren Creek ll-Vni-60 (AOS). 

80, Plebejus (Icaricia) acmon (West. & Hew.) Reserve 14-VII-33 (JSG); 
pShonT“ 25^ra33 "(JS^ Kerrick 27-VII-34 (EDG); Coldwater 3- 
Vm-34 (EDG); Smoky Jack 4-Vn-54 (JWT); Mather lltol3-VII- 

56 (JSG); Hetch-Hetchy Stimmit 13-VII-56 (JSG); Gin Flat IT'-VIII- 

57 (JWT); Tenaya Canyon ll-VII-SS (AOS); Tuolumne Meadows 3- 

.IX-58 (JWT); Jerseydale 30-V-59, 27-V-60 and ll-VI-61 (JWT), 

26- VI-59 (JSG); Indian Flat 30-V-59, 27-V-60and 14-V to 10-VI- 

61 (JWT); Briceburg 21-111-61 (KT); El Portal 21-111-61 (KT), 14- 
V-61 (JWT); Cliff House 9-VI-61 (JWT); Mather 10- VI-61 (JWT); 
Cathay ll-VI-61 (JWT). Mono Lake 30- VI-41 (JSG), 30- VI to 7- 
Vn-58 and 12- VI- 62 (AOS); Huntoon Public Camp, Bridgeport 19- VI- 
54 (JSG); Warren Creek ll-VIII-60 and 26- VI-61 (AOS). 

81, Plebejus (Icaricia) lupini (Bdv. ) Eagle Peak l-Vn-33 (JSG); Dana, 

W slope l6^VnL52T'T^VIII-53 and 20-Vn-58 (JWT); Smoky Jack 3 
to 4- VII- 54 (JWT); Crest W of Tioga Pass 12-VI1-57, 4-Vn to 19- 
Vni-58, 15 to 23-Vn-60 and 25-VI-61 (AOS); Upper Gaylor Lake 14- 

to 19-VII-58 (AOS); Helen Lake 18- VII- 58 (AOS); Dana, N slope 24- 
Vm-58 (AOS), Lee Vining 3 to 5-Vn-38 (JWT); E above Saddlebag 
Lake 28-VII-58 (AOS); Below Conway Summit 23-VI-62 (JWT). 

82, Everes comyntas (Godt. ) Mono Lake 5-Vn-38, 30- VI- 50 and 23- VL 
6TTJW T)r~l6^I- 6 1 and 12- VI- 62 (AOS); E above Saddlebag Lake 
28-VII-58 (AOS), 

83, Everes amyntula Bdv. Hetch- Hetchy Siimmit 13-VU-56 (JSG, seen); 
Jerseydale 30^-59 (JWT). 

84, Philotes enoptes (Bdv.) Yosemite Valley 17-VI-32 (JWT); Ledg-e 9- 
VIL33 IjSOF^eserve l6-Vn-33 and 7-Vn-56 (JSG); Tioga Pass 
16-VIL52 (JWT), 12-Vn-57, 9-Vn to 14-Vni-58 and 14 to 23-Vn-60 
(AOS); Smoky Jack 4- VII- 54 (JWT); Big Oak Flat Road 5- VII- 54 
(JWT); Tenaya Canyon ll-VII-58 (AOS). E above Saddlebag Lake 
28- ¥11- 58 (AOS). 

85a. Philotes battoides battoides (Behr) Reserve 15 to 16-VII~33 and 7- 

Vn-56 (JSG);“’^dr'c?^‘^^ 29-Vn-34 (EDG); Gaylor Lakes Trail 16- 
Vni-52, 6-Vni-53 and 3.IX-58 (JWT); Mt. Hoffman 14.Vn-57 (AOS); 
Pilot Peak 18- Vn-57 and 3 1- Vin-58 (JWT); Crest W of Tioga Pass 
19~Vn to l-Ym-58, 14 lo 15-Vn-60, 25- VI- 61 and 8- Vm- 62 (AOS); 
Helen Lake 18to29“Vn-58 (AOS); Cockscomb Peak 26-VII-58 (AOS); 
Vogelsang 23-Vn-58 (JWT); Vogelsang Pass 3- ¥111-58 (AOS), 
b. Philotes battoides glaucon (Bdv.) Mono Lake 5- VII- 38 and 23- VI- 62 
(JWT), 30-YI-41 NSGfr^ 30- VI-58 and 26 to 27-VI-61 (AOS); War- 
ren Creek 21-Vn~58 (AOS); Conway Summit 23-VI-62 (JWT). 

86, Philotes speciosa (Hy, Edw. ) Mariposa 30-V-32 (G. & Bohart); 

(R. M, Bohart), 

87, Phaedrotes piasus (Bdv.) Eagle Peak l-Vn-33 (JSG); Reserve 9 to 
TATvn^i^lED^; Jerseydale 9 to 13- VI- 62 (AOS); Mather 30- YI- 

62 (JWT). Mono Lake 30-VI-41 (JSG). 


2( i ): i -96, 1963 


YOSEMITE BUTTERFLIES 


87 


88a. Glaucopsyche lygdamus behrii(Edw.) Indian Flat 16 to 17-IV-61 (JWT); 
El Porting- IV- 6F7 JwT)7“ 

b. Glaucopsyche lygdamus Columbia (Skin.) Eagle Peak l-VII-33 (JSG); 
Reserve 9 to 14- VII-34 “[eDG ?); Tioga Pass 16- VIII- 52 (JWT); 

Crane Flat and Tamarack Flat 3- VII- 54 (JWT); White Wolf 10- VII- 
56 (JSG); Tenaya Canyon ll-Vn-58 (AOS); Upper Gaylor Lake 19- 
VII-58 (JWT); Dana, W slope 20-VII-58 (JWT); Below Vogelsang 
Pass 3-VIIL58 (AOS); Badger Pass 23-VI-59 (JSG). E above 
Saddlebag Lake 28- VII- 58 (AOS). 

89 . Celastrina argiolus echo (Edw. ) El Portal 28- VI-33 (JSG); Glacier 
' 9 - VII- 33 (JSG); ReT^e 14 to 18- Vn-33 and 7- VII-56 (JSG); Crane 
Flat 3-VII-54 (JWT); Mather 12-Vn-56 (JSG), 14-V-61 (JWT); 
Aspen Valley 14- VII- 56 (JSG); Darrah, Jerseydale, and Indian Flat 
29 to 30-V-59 and 10- VI-61 (JWT); Badger Pass and Bridal Veil 23- 
VI - 59 (JSG); Tamarack 24- VI- 59 (JSG); Briceburg and El Portal 
21-III-61 (KT). 

PIERIDAE 

90a. Anthocharis (Anthocharis) sara sara Luc. Indian Flat 13-V-61 (JWT); 

El Portal 14-V-61 (Jl^). “ “ 

Anthocharis (Anthocharis) sara reakirtii Edw. , gen. vern. Brice- 
burg 21-III-6T~(W), 16- VI- 61 (JWT); El PortST" 21-III and 4-IV- 

61 (KT), 14-IV-62 (JWT); Indian Flat 17-IV-61 (JWT). 

b. Anthocharis (Anthocharis) sara stella Edw. 'Eagle Peak l-VII-33 
*^S^ ReserT^ 16 to 17- VII-33" ( JSG);"'"- White-'Wolf lO-Vn-56 (JSG). 

91 . Anthocharis (Falcapica) lanceolata Luc. Eagle Peak l-VII-33 (JSG); 
Ledge 9 - VII- 3 3 (JSG); Tosemltr Valley 14-V-59 (R. Patil Allen); 
Jerseydale 9 to 13-VI-62 (AOS). 

92 . Euchloe creusa hyantis (Edw.) Eagle Peak l-VII-33 (JSG); Glacier 
9^VILT3 (JSG); Reserve l6-Vn-33 (JSG); Ostrander 4- VII- 34 
(EDG); El Portal 16- IV and 14- V- 6 1, andl4-IV-62 (JWT, common); 
Indian Flat 17-IV-61 (JWT); Mather 14-V-61 (JWT). 

93 . Euchloe ausonides coloradensis (Hy. Edw.) Reserve 16 and 17-VII- 
33 TJSG); Tamara^ Flat 3- VII- 54 (JWT). Mont) Lake 30- VI- 58 
(AOS). 

94 . Colias eurytheme Bdv. Ledge 9- VII- 33 (JSG); Reserve l6-Vn-33 

and 7 to“TlTviI-56 (JSG); Camp 9, Yosemite Valley 23-Vn-33 (JSG); 
Crane Flat 3-VII-54, 18-Vm-57, and 10- VI-61 (JWT); Mather 11 

to 12-VII-56 (JSG), 14- V and 10- VI- 61 (JWT); Hetch-Hetchy 13-VII- 

56 (JSG); Aspen Valley 14- VII- 56 (JSG); Gaylor Lakes Trail 19- VII- 

58 (JWT); Dana, W slope 30-VIII-58 (JWT); Briceburg 21-III-61 
(KT); El Portal 21-III-61 (KT), 14.V-61 (JWT); Yosemite Valley 

I 9 -IV- 6 I (KT); Indian Flat 13-V-61 (JWT); Jerseydale II-VI- 6 I 
(JWT). 

95 . Colias philodice hagenii Edw. Mono Lake 30-VI-41 (JSG); 18-VII- 

57 and 30-VI-58 "JKosY. 

96 . Colias occidentalis chrysomelas Hy. Edw. "Yosemite" VI-26 (E. 

O. Essig). 

Colias behrii Edw. Tioga Pass 13-Vni-29 and 1 to 3-Vin-56 (JSG), 
14-VII-31, 13-Vin-50 and l6-Vni-57 (JWT), 4-Vni-57 and 14-Vnto 
19-Vni-58 (AOS); Florence 5-Vni-33 (JSG); Kuna 7-Vin-33 (JSG); 
Kerrick 27-VII-34 (EDG); Snow Lake 29-Vn-34 (EDG); Tuolumne 


88 


GARTH AND TILDEN 


/. Res. Lepid. 


Meadows 15-Vni-57 (JWT); Pilot Peak 15 to 18- VIU- 57 (JWT); 

Upper Gaylor Lake 19-VII to 20-Vni-58 (AOS); Helen Lake 29-VII- 
58 (AOS); Crest W of Tioga Pass 31-VII-58 (AOS); Rafferty Creek 
Trail 3-Vni-58 (AOS, JSG); Merced Lake Trail 4-VIII-58 (AOS); 

Lyell Base Camp 10- VIII- 58 (AOS); Elizabeth Lake 23- VIII- 58 (AOS). 

E of Donohue Pass 4 to 6-VIII-59 (AOS), 

98. Colias (Zerene) eurydice Bdv. "Yosemite" VI-26 (E, O. Essig or 

PTJTWooIfJT"^ 

99. Nathalis iole Bdv. "Yosemite" VI-26 (P. J. Woolf). Warren Creek 
20, 21 ai^n8-VU-58 (AOS). 

100. Neophasia menapia tau (Scud.) Big Oak Flat Road VIII-28 (JSG, seen); 
Little Yosemite ValT^ 3- VIII- 33 (JSG); Pate 23- VII- 34 (EDG); 

Buck Meadow 4-VII-54 (JWT); Unicorn Peak 27-VII-59 (AOS). Dead- 
man's Summit 20-IX-56 (JWT); June Lake 2 1-VIII-57 (JSG), 

101. Pieris (Pontia) beckeri Edw. Mono Lake 5-VII-38, 30-VI-50, 15- 

Vni-53 and 23 -VI- 62 (JWT), 26 and 27-VI-61 (AOS). 

102. Pieris (Pontia) sisymbrii Bdv. Eagle Peak l-VII-33 (JSG); Reserve 
16- VII- 33 and 7-Vn-56 (JSG); Dana, W slope 19"Vn-52, 16-VIII-53 
and 20-Vn-58 (JWT); Upper Gaylor Lake 4 to 8-VII-57, 9 to 19-Vn- 
58 and 25-VI-61 (AOS); Crest W of Tioga Pass 9-VII-58 (AOS); 

Tioga Pass lO-Vn-58 (AOS); Briceburg and El Portal 2 l-ni-6 1 (KT); 

El Portal 14-IV-62 (JWT). 

103. Pieris (Pontia) protodice Bdv. h Lee. El Portal 28-VII-33 (JSG); 
Yosemite Creek Trail 9 to 17-Vn-56 (JSG). 

104a. Pieris (Pontia) occidentalis occidentalis Reak. Crest W of Tioga Pass 
181^20- Vm- 57 and 19 to 2”0-VIII-58 (AOS), 
b. Pieris (Pontia) occidentalis calyce Edw. Pilot Peak 18- VIII- 52, 15- 
Vni-57, and“20-VII-58 (JWT); Crest W of Tioga Pass 8-VII-57 and 
9-VII-58 (AOS); Upper Gaylor Lake 9-VII-58 (AOS), l-Vm-58 
(JSG); Vogelsang Camp 3-VIII-58 (JSG). Warren Creek 29-VIII-57 
(AOS). 

105. Pieris (Pieris) napi venosa Scud. Yosemite Valley 14-V-59 (R. P. 
Allen); Briceburg 21-III-61 (KT); El Portal 14-IV-61 (JWT); Jer- 
seydale 22-VI-61 (AOS). 

106. Pieris (Pieris) rapae (Linn.) Pate 23-VII-34 (EDG); Indian Flat 30- 
V-59 (JWT, se^l7“ 

PAPILIONIDAE 

107. Papilio zelicaon Luc. Reserve l6-Vn-33 (JSG); Mather 15-VII-56 
(JSG); Jerseydale 16- VI- 57 and 4- VUI- 62 (AOS); Crest W of Tioga 
Pass 9-VII-58 (AOS); Upper Gaylor Lake 19-VII-58 (AOS); El Por- 
tal 4-IV-61 (KT). 

108. Papilio indra Reak, Yosemite Valley 17-VI-32 (JWT); Reserve 18- 

VII- 3r (A. Carthew); Crest W. of Tioga Pass 25- VI-61 (AOS). 

Mono Lake 23-VI-62 (JWT). Mammoth Peak 16-VII-60 (AOS). 

109. Papilio rutulus Luc. Camp 19, Yosemite Valley 26- VI-33 (JSG); 

Pate 23- VII- 34 (EDG); Yosemite Creek Trail 9-Vn-56 (JSG); 
Hetch-Hetchy 13-VII-56 (JSG); Darrah, Jerseydale, and Indian Flat 
29 to 30-V-59 (JWT). Mono Lake 30- VI-41 (JSG, seen), 27-VI-61 
and 12- VI- 62 (AOS); W of Lee Vining 27-VI-61 (AOS); Below Conway 
Summit 23- VI-62 (JWT). 

Papilio multicaudatus Kby. Jerseydale 7 -VIII- 5 6 and 2- VIII- 57 (AOS). 


2(i)ti-s6, 196} 


YOSEMITE BUTTERFLIES 


89 


Ilia. Papilio eurymedon eurymedon Luc. Ledge 9-VII-33 (JSG); Pate 23- 
Vn- 34 " (EDG); Mather 12-^-56 (JSG); Hetch-Hetchy 13-Vn-56 
(JSG); Jerseydale 16- VI- 57, 16- VI-58, 21-VI-61 and 13-VI-62 (AOS); 

Darrah, Jerseydale, and Indian Flat 29 to 30-V-59 (JWT). 
b. Papilio eurymedon albanus F. & F. Research Reserve 10- VII- 56 

IjSGir 

112. Parnassius clodius baldur Edw. Tioga Pass 7-VII-31 (JWT); Eagle 

Peak 1-VII^3 (JSG)r^lacier 9- VII- 33 (JSG); Dana 8- Vni-33 ( JSG); 
Slide 29-VIL34 (EDG); Gin Flat 4-Vn-54 (JWT, seen); Jerseydale 
23-VI-56, 21-VI-57. 16 to 18- VI- 58 and 22- VI- 6 1 (AOS); Yosemite 

Creek 7-Vn-56 (JSG); White Wolf 10- VII- 56 (JSG); Crest W of 
Tioga Pass 12-VII to IS-VIH-ST, 28- VII to 19-VIII-58 and 4 to 20-VII- 
60 (AOS); Pilot Peak 18- VII- 57 and 31-Vin-58 (JWT); Rafferty Creek 
21- VII- 57 (AOS); Tenaya Canyon ll-VII-58 (AOS); Lewis Creek 
Trail 3-Vni-58 (JSG); Dana, W slope 30-VIII-58 (JW-T). 

113. Parnassius phoebus behrii Edw. 

Reserve 14- VH^S' (“180)7 Dana8-Vm-33 (JSG); Pilot Peak 13 to 
14- VIII- 52, 18-Vn to 15-Vni-57 a-nd 31-VIII-58 (JWT); Upper Gaylor 
Lake 3 1- VII to 20- VHI- 57 and 3 1- VII to 20- VIU- 58 (AOS), 19-VII-58 
(JWT), l-Vin-58 (JSG); Helen Lake 18- VII- 58 (AOS); Dana, W 
slope 20-VII-58 (JWT); Crest W of Tioga Pass 19-VIII-58 (AOS); 
Dana, N slope 24- VIE- 58 (AOS). E of Saddlebag Lake 28-VII-58 
(AOS). . 

114. Epargyreus clarus (Cram j Coulterville Road VI-26 (D. D. MacLean); 
Indian Flat 30-V-59 (JWT, seen). 

115. Thorybes pylades (Scud.) Fish Camp 18- VI- 57 (JWT); Jerseydale 
JO^V^T^ (“jWrjT^Indian Flat 13-V-61 (JWT). 

116. Thorybes mexicana nevada Scud. Tioga Pass 7-VII-31, l6-Vni-52 

and 18-VIL5TnJ^Tir~8TviI-57, 26- VI to 19- VH- 58, 14-VII-60 and 

25-VI-61 (AOS); Reserve 16-VII-33 (JSG); Tamarack 3-Vn-54 
(JWT); Pilot Peak 18-VII-57 and 3 1- VIE- 58 (JWT); Dana, W slope 
I 7 .VEI -57 and 20-VE-58 (JWT); Gaylor Lakes Trail 9-VII-58 (JWT); 
Crest W of Tioga Pass 9-VE-58 (AOS); Upper Gaylor Lake 14 to 
19-VII-58 (AOS), l-VIE-58 (JSG); Helen Lake 18 to 29-VII-58 (AOS); 
Gin Flat and Crane Flat lO-VI-61 (JWT). 

117. Thorybes diversus Bell Aspen 14-VE-56 (JSG); Jerseydale I 6 -VI- 
5Ta^n6-"vi-58 fAOS); Mather 10- VI- 61 and 30- VI- 62 (JWT). 

118. Pyrgus ruralis (Bdv. ) Reserve 15- VE- 33 and 10- VII- 56 (JSG); Tam- 
arack 3-VII-54 (JWT); Tenaya ll-VII-58 (AOS, seen); Lyell Fork 
Meadows 7- VIE- 58 (AOS). 

119 . Pyrgus communis (Grote) "Yosemite" VI-26 (P, J. Woolf); Aspen 

' Valli^ 14-VE-56" (JSG); El Portal 16-IV, 14-V and lO-VI-61 (JWT); 

Indian Flat 15-IV, 14- V and 1 1- VI-61 (JWT), 

120. Heliopetes ericetorum (Bdv.) Jerseydale 16-VI-58 (AOS); El Portal 

14- V-61 (JWT, s7^n)7 Highway 120 and Hetch-Hetchy Road 14-V- 61 
(JWT). 

121. Erynnis persius (Scud.) Reserve 15-VII-33 (JSG, as afranius); Chow- 
chilla Mt. 26-VI-54 (JWT); Aspen Valley 14-VII-56 (JSG); Mather 

15- VE-56 (JSG); Badger Pass 23-VI-59 (JSG); Indian Flat 17 -IV- 6 1 
(JWT); Jerseydale 21to23-VI-6l (AOS). 

122. Erynnis pacuvius lilius (Dyar) Chowchilla Mt. 26- VI- 54 (JWT, com- 
mon); Tamarack fET™ 3 - VII- 54 (JWT); Crane Flat 24- VI- 59 (JSG). 
Erynnis propertius (Scud. & Burg.) Ledge 9-VII-33 (JSG); Reserve 


90 


GARTH AND TILDEN 


/. Res. Lepid. 


16-VII-33 (JSG); Crane Flat 3-Vn-54 (JWT); Yosemite Creek 9-VII- 

56 (JSG); Mather 12-VII~56 (JSG), 10- VI- 61 (JWT); Hetch-Hetchy 

Sixmmit 13-Vn-56 (JSG); Badger Pass 23-VI-59 (JSG); Briceburg 
21-in-6l (KT); El Portal 21-ni-6l (KT), 14-V-61 (JWT); Yose- 

mite Valley 19-IV-61 (KT); Indian Flat 13-V and 10- VI-61 (JWT); 
Jerseydale 17- VI- 60, 2 1- VI- 6 1 and 4 to 13- VI- 62 (AOS). 

Erynnis zarucco fnneralis (Scud. & Burg.) Coulterville Road VI-26 
(D. D. MacLean). 

125. Erynnis tristis (Bdv. ) No Yoserhite record. 

126a. Hesperia harpalus harpalus (Edw. ) Mono Lake 17-VII to 26-Vni-57, 
30-VI to 20-VII-58 and 26-VI-61 (AOS), 26-Vin-57 and 20- VH- 58 
(JWT); Bodie 18-VII-57 (AOS); Lee Vining 24- VI-58 (AOS); War- 
ren Creek 26-VI-61 (AOS). 

Hesperia harpalus (Edw.), "blend zone" form Eagle Peak l-VII-33 
(JSG); Reserve 16- VII- 33 (JSG); Yosemite Creek 9 to 17-VII-56 
(JSG); Rafferty Creek 21-Vn-57, 24- Vn to 3- Vin-58 (AOS); Dana, 

W slope 16-VIII-57 and 30-VIII-58 (JWT); Crest W of Tioga Pass 18 
to 20-VIII-57, 20-Vin-58 and 20 to 28-Vn-60 (AOS); Merc.ed Lake 
Trail 4-VIII-58 (AOS); Unicorn Peak 12- VIII- 58 and 27- VB- 59 (AOS); 
Gaylor Lakes Trail and Pilot Peak 31-VIII-58 (JWT); Tuolumne 
Meadows 3-IX-58 (JWT); Cold Canyon 25-Vn-60 (AOS); Dana, N 
slope 8-Vin-60 (AOS). 

b. Hesperia harpalus yosemite Leuss. Big Oak Flat Road, near entrance 
to Tuolumne Grove 2- IX- 29 (J. Strohbeen and A. E. Dodge, the type 
series); Carl Inn ll-VII-31 (JWT); Wawona 27-Vm-46 (JWT); 

Smoky Jack 4-VII-54 (JWT); Jerseydale l-Vin-57 (AOS). 

127. Hesperia miriamae MacNeill Mono Pass, Inyo County VIII- 56, VIII- 

57 and Vni-58 (Don MacNeill, the type series); Unicorn Peak 26-Vn, 

12 and 23- VIII-58, and 27-Vn-59 (AOS); Cockscomb Peak 26-Vn-58 
(AOS); Dana, N slope 31- VII- 58 and 8-Vin-60 (AOS). 

128. Hesperia nevada (Scud.) Mono Lake 26-VI-61 (AOS); Warren Creek 
26- VI- 61 and 12"-VI-62 (AOS). 

129. Hesperia juba (Scud.) "Yosemite" VI-26 (collector unknown); Old 
Village lTrvj-32 (McQuesten); Mather 10- VI-61 (JWT). Mono 
Lake 27- VI-61 (AOS). 

130. Ochlodes sylvanoides (Bdv.) Yosemite Valley Vn-33 (JSG); Pate 
Van^7~23”vil- 34 TeDG); Hetch-Hetchy 13- VII- 56 (JSG); Crane 
Flat 18- VIII- 57 (JWT); Mather 4-IX-57 (JWT); Jerseydale 2-IX- 

58 (AOS). Mono Lake 14 and 15-Vm-51 and 13 to 15-VIII-52 (JWT). 

13 1. Ochlodes agricola (Bdv. ) Darrah 29-V-59 (JWT); Indian Flat 30-V- 

59 (JWT); El Portal 14-V-61 (JWT). 

132a. Polites sabxileti sabuleti (Bdv.) Bridgeport 14-Vni-50 and 16-Vni-51 
(JWT); Mono La'ki^2“3rvi- 62 (JWT). 
b. Polites sabuleti tecumseh Grin. Tioga Pass 7-Vn-31, 13-VIII-50, 

Ts^^^-TITTs^VLElSZ and 18-Vnto 16- VIII- 57 (JWT), 14-VIIto7- 
VIII- 58 (AOS); Reserve 17-Vn-33 and 10- VII- 56 (JSG); Kuna 7-Vm- 
33 (JSG); Kerrick 27-VII-34 (EDG); Snow Lake 29-VII-34 (EDG); 
Tuolumne Meadows 15-VIII-57 and 3-IX-58 (JWT), 15-Vin-58 (AOS); 
Gin Flat 17-Vn-57 (JWT); Rafferty Creek 24-VII-58 (AOS), 3-Vin- 
58 (JSG); Upper Gaylor Lake l-Vin-58 (JSG); Upper Lyell Base 
Camp 9-Vin-58 (AOS); Elizabeth Lak% 12-Vin-58 (AOS); Dana, W 
slope 30-Vni-58 (JWT); Pilot Peak 31-Vin-58 (JWT). 


2(t)tl-96, 196} 


YOSEMITE BUTTERFLIES 


91 


133. Polites sonora (Scud, ) Ledge 9-VII-33 (JSG); Reserve 17-Vn-33 

(JSG); Tioga Pass l6-Vni-52 (JWT); Aspen Valley 14-Vn-56 (JSG); 
Crane Flat 18 and 19"Vn-57 (JWT); Gin Flat 19-VII-57 (JWT); 
Tuoltimne Meadows 14-Vin-57 and 3-IX-58 (JWT), 15 to 21-VIII-58 
(AOS); Bridal Veil 23-VI-59 (JSG); Mather 10- VI- 6 1 and 30- VI- 62 
(JWT). Mono Lake 4-VII-33, 5-VII-38, 30-VI-50 to IS-VIII-SO and 

15-Vni-51 (JWT), 15-Vni-57, 30-VI-58 and 27-VI-61 (AOS); Bodie 

18- VII- 57 (AOS); Warren Creek ll-VIII-60 (AOS); W of Lee Vining 
26-VI-61 (AOS). 

134. Amblyscirtes vialis (Edw.) Yosemite Valley, Meadow near Ahwahnee 
HStirT7^VL3T'‘7jWT); Jerseydale 17- VI- 57, 22- VI- 6 1 and 9 to 13- 
VI- 62 (AOS). 



YOSEMITE PARK 

1933 Localities: 1 — Camp 19; 2 — ^Camp 9; 3 — Mt. Dana; 4 — Eagle Pk.; 
5 — El Portal; 6 — Florence L.; 7- — Glacier Pt.; 8 — Glen Aulin; 9 — 'Mt. 
Kuna; 10 — Ledge Tr.; 11 — Little Yosemite; 12 — Mt. Lyell; 13 — Mu- 
seum; 14 — Pohono Tr.; 15 — Research Reserve; 16 — Wawona. 

1934 Localities: 17 — Benson L.; 18- — Coldwater Can.; 19 — KerrickPass; 
20 — Ostrander L.; 21 — Pate Valley; 22 — Slide Can.; 23 — Tenaya L. 

Key 


Auto Rd. 


- - Trails Covered 1933 .... Trails Covered 1934 


92 


GARTH AND TILDEN 


/. Res. Lepid. 


APPENDIX III 
YOSEMITE LOCALITIES 

West- slope localities visited by John S. Garth in 1933 (See Map 1): 

1. Camp 19; Yosemite School of Field Natural History headquarters near 
Sentinel Bridge, floor of Yosemite Valley (3960). Transition Zone; 
Meadow Association. 

2. Camp 9: Meadow near Camp Curry in Yosemite Valley (4000), a fav- 
orite haunt of Argynnis cybele leto. Transition Zone; Black Cotton- 
wood; Meadow Association. 

3. Dana: Trail from Tuolumne Meadows (8600) to top of Mt. Dana (13, 050). 
Hudsonian, Arctic-Alpine Zone; Subalpine Forest; Alpine Fell-Fields. 

4. Eagle: Trail from top of Yosemite Falls (6525) to summit of Eagle Pk. 
Canadian Zone; Red Fir Forest; Moist Meadow; Riparian Association. 

5. El Portal: Highway 140 near El Portal (1919) on the Merced River; short 
distance up Coulterville Road. Upper Sonoran Zone; Chaparral; Stream. 

6. Florence: Lake above Lewis Creek Trail (10,500). Hudsonian Zone, in 
close proximity to Arctic- Alpine. Subalpine Forest; Montane Meadows. 

7. Glacier Trail from Glacier Point (7214) to top of Nevada Falls (5910). 
Canadian Fir Forest replaced by Upper Sonoran Chaparral after fire. 

8. Glen Aulin: High Sierra Camp on Tuolumne River at confluence of Con- 
ness Creek (7850). Canadian Zone; Transition elements (A. hydaspe). 

9. Kuna; From Lyell Base Camp (10, 600) to summit of Kuna Crest (12, 000), 
down Dana Fork to Tuolumne Meadows (8600). Hudsonian, Arctic-Al- 
pine Zone; cirques, tarns, snow banks, talus slopes, alpine meadows. 

10. Ledge: Short trail from Camp Curry (4000) to Glacier Point (7214). 
Transition, Canadian Zone; steep cliffs, talus slopes, springs, stream, 

11. Little Yosemite: Hanging Valley above Nevada Falls (5910) on Merced 
Lake Trail. Transition Zone; Jeffrey Pine, White Fir, Western Juniper. 

12. Lyell: From Vogelsang Lake (10, 500) to summit of Mt. Lyell (13, 090). 
Hudsonian, Arctic- Alpine Zone; glaciers, tarns, Alpine rock gardens. 

13. Museum: Garden along diverted stream behind Yosemite Museum (4000). 

14. Pohono: Trail from Glacier Point (7214) over Sentinel Dome (8117) to In- 
spiration Point (5391). Canadian and Transition Zone; stream, meadow. 

15. Reserve; Boundary Hill Research Reserve, a 25- square-mile area bound- 
ed by the Tioga Road, Yosemite Creek, Cascade Creek, and the north 
rim of Yosemite Valley. (6525 - 9200). Canadian Zone with weak Hud- 
sonian atop Research Ridge; Red Fir Forest; Lodgepole Pine; Western 
Juniper. 

16. Wawona: Highway 41 (Wawona Road) from Chinquapin (6266) to Wawona 
(4096). Transition Zone; Dry Hillside; Monardella and Erysimum . 

West- slope localities visited by Edmund D, Godwin in 1934 (See Map 1): 

17. Benson: Trail from Pate Valley (4500) over Pleasant Ridge (8000) to 
Pleasant Valley (7000) and Benson Lake (8000). Transition, Canadian 
Zone; Upper Sonoran influence ( Cercyonis silvestris, Coenonympha). 

18. Coldwater : Trail from Virginia Creek (8500) to Glen Aulin (7800) along 
Coldwater Canyon. Canadian Zone; Red Fir Forest; Riparian Assoc. 

19. Kerrick : Trail from Benson Lake (8000) through Seavey Pass (9100) to 
Kerrick Canyon (8900), thence to Kerrick Meadows (9400). Canadian 
to Hudsonian Zone; Red Fir, Lodgepole Pine Forest; Montane Meadow. 


2(i)xi-96, 196} 


YOSEMITE BUTTERFLIES 


93 


20. Ostrander: Trail from Glacier Point Road (7100) to Ostrander Lake 
(8500). Canadian to Hudsonian Zone border ( Tsuga mertensiana). 

21. Pate: Trail from Harden Lake (7575) down north- facing granite slope 
to Pate Valley (4500). Canadian to Transition Zone; Upper Sonoran 
influence from opposite exposure. Monardella and Umbellularia. 

22. Slide: Trail from Kerrick Meadows (9400) to Snow Lake (10, 200) to 
Slide Canyon (10, 000). Hudsonian Zone; Subalpine Meadow Associa- 
tion (Colias behrii ). 

23. Tenaya: McGee Lake Trail from Glen Aulin (7850) on the Tuoltimne 
River to Tenaya Lake (8141). Canadian Zone; Red Fir; Lodgepole Pine. 

Western Foothill localities visited from 1956 to 1962, mostly outside 

Yosemite National Park: 

Bear Creek Lodge : On Highway 140 between Midpines (2477) and Brice- 
burg(1186). Transition Zone; Yellow Pine Forest; Riparian Association. 

25. Big Oak Flat Road : Highway 120, the part between Groveland (2846) and 
Carlon (4624) within Stanislaus N F. Upper Sonoran, Transition Zone. 

26. Briceburg: Mariposa Co. , on Highway 140 in Merced River Bottom 
(1186). Upper Sonoran Zone; Chaparral and Coastal Sage Scrub. 

27. Buck Meadows: Stanislaus N F, on Highway 120 (3006). Transition Zone; 
Yellow Pine Forest with "Mountain Misery" ( Chamaebatia ) understory. 

28. Carl Inn: Stanislaus N F, at juncture of Hetch-Hetchy Road to Mather 
and Old Tioga Road (4718). Transition Zone; Mixed Forest, cut over. 

29 . Cathay: Mariposa Co., on Highway 140; Upper Sonoran; Oak Woodland. 

30. Chowchilla Mtn. : Sierra N F, W of Fish Camp (4982) on Chowchilla Mtn. 
Road, near Bear Wallow turnoff. Canadian Zone; Red Fir Forest. 

31. Cliff House : Stanislaus N F, on Highway 120 N Fork, Tuolxxmne River, 
crossing. Transition Zone; Yellow Pine Forest; Y erba Santa. 

32. Darrah: Mariposa Co. (3400). Upper Sonoran, Transition Zone; Chap- 
arral; Coastal Sage Scrub; Yellow Pine, Mixed Forest; Riparian Assoc. 

33. El Portal: Yosemite N P, on Highway 140 in Merced River Bottom (1919). 
Upper Sonoran Zone; Chaparral, Coastal Sage Scrub; Riparian Assoc. 

34. Groveland; Stanislaus N F, on Highway 120 (2846). Transition and Upper 
Sonoran Zone; Yellow Pine Forest; Mixed Forest; Chaparral. 

Highway 120 and Hetch- Hetchy Road : Juncture, Big Oak Flat Road and 
Mather Road (4624). Upper Sonoran, Transition; Oak Wood; Yellow Pine. 

36. Indian Flat; Sierra N F, on Highway 140 in the Merced River bottom 
(1553). Upper Sonoran Zone; Coastal Sage Scrub, Chaparral; Yellow Pine. 

37. Jer seydale: Sierra N F, (3800). Transition Zone; Yellow Pine, Mixed 
Forest; Chamaebatia, Monardella ( Argynnis hydaspe); some Meadowland. 

38. Mariposa: Mariposa Co. , on Highway 140 (2046). Upper Sonoran, Transi- 
tion Zone; Chaparral, Coastal Sage Scrub; Mixed Forest; Riparian Assoc. 
Mather: Stanislaus N F (4522), on Hetch-Hetchy Road. Transition Zone; 
Yellow Pine, Mixed and White Fir Forest; Montane Meadow; Riparian. 
Includes Carnegie Experimental Garden (Hog Ranch) 1 mi NE of Mather. 

40. Oakhurst: Madera Co., on Highway 41. Upper Sonoran Zone; Foothill 
Woodland; Oak and Digger Pine with Ceanothus and grass (Coenonympha). 

West-Slope Mid- Elevation Localities visited from 1956 to 1962, all within 
Yosemite National Park: 

41. Aspen Valley : On Old Tioga Road above turnoff from Mather Road at Carl 
Inn ( 639 O). Canadian Zone; Red Fir Forest; Montane Meadow; lupine. 


94 


GARTH AND TILDEN 


/. Res. Lepid. 


42. Badger Pass: Meadow beside Badger Pass Ski Lodge on Glacier Point 
Road (7300). Canadian Zone; Red Fir Forest; Wet Meadow Association. 

43. Bridal Veil : Public Camp on Bridal Veil Creek where crossed by Glac- 
ier Point Road (7000). Canadian Zone; Red Fir Forest; Riparian Assoc. 

44. Crane Flat: On Highway 120 where it becomes the Tioga Road (6192). 
Transition Zone; Mixed Forest; Montane Meadow. Also roadside above. 

45. Gin Flat: On Tioga Road above Crane Flat (7036), Canadian Zone; 

Montane Meadow. 

46. Glacier Point: Road to, with maximum elevation 7800 feet, Canadian 
Zone; openings in Red Fir Forest; Montane Meadow; Riparian Assoc. 

47. Hetch- Hetchy : Road from Mather (4522) to Hetch-Hetchy Reservoir 
(3796). Upper Sonoran Zone; Digger Pine; Manzanita. Summit (5027) 
Transition Zone; White Fir Forest; Streamside Association; lupine. 

48. Mirror Lake: Floor of Yosemite Valley (4082). Transition Zone; Black 
Oak, White Fir, Incense Cedar, Cottonwood, Ceanothus, Coffee Berry. 

49. Old Village: Yosemite Valley floor (3964). Transition Zone; Riparian. 

50. Research Reserve: Meadow in Blue Jay Creek (7900). Canadian Zone; 

Red Fir, Lodgepole Pine. Research Ridge (8700) Dry Hillside Assoc. 

51. Smoky Jack; Public Camp on Tioga Road, above Gin Flat. Canadian 
Zone; Red Fir Forest; Montane Chaparral. 

52. Tamarack Flat: Meadow above Tamarack Public Camp on Tioga Road 
(6390). Upper Canadian Zone; Red Fir; Lodgepole Pine; Ceanothus. 

53. Tioga Road; North-facing slope 1 mi W of White Wolf turnoff (8000) and 
first summit beyond White Wolf (8090). Canadian Zone; Red Fir Forest. 

54. Tuolumne Grove : On Highway 120 W of Crane Flat (5500-6000). Tran- 
sition Zone; Mixed Forest; Big Tree; Riparian Assoc. Polygonia zephyrus. 

55. White Wolf; One mi N of Tioga Road (8090). Canadian Zone; Red Fir 
Forest; Montane Meadow Assoc. Plebejus saepiolus, P. glandon podarce. 

56. Wawona : Open forest between Wawona Lodge (4096) and the turnoff to 
Giant Forest (4350). Transition Zone; Yellow Pine Forest. 

57. Yosemite Creek: Trail S from Public Camp on Tioga Road (7200). Can- 
adian Zone; Riparian and Wet Meadow; Western Juniper on drier slopes. 

58. Yosemite Valley: Floor of Valley (4000). Transition Zone; Mixed Forest; 
Riparian Association. Includes Meadow near Ahwahnee Hotel. A. vialis. 

High Country localities visited since 1956 (those near Tioga Pass since 
1929): 

59. Agnew Pass: Inyo N F (9946). Hudsonian Zone; open sage slope. 

60. Bert Lake : N slope of Mt. Maclure (11, 700); Arctic- Alpine; rock gardens. 

61. Cockscomb Peak: Cathedral Range (11,000); Arctic-Alpine; White-Bark 
Pine; gradual slope; Spraguea in gravelly areas. Hesperia miriamae. 

Dana , N slope: (11,000- 13,050 ). Arctic-Alpine; rockslides, flowers, 
grass. Lycaena p. hypophlaeas . Dana, W slope: From Tioga Pass (9941) 
to timber line (11, 000). Hudsonian to Arctic- Alpine; Subalpine Forest; 
Alpine Fell-Fields. 

Dog Lake Trail: From Tuolumne Meadows ( 8600 ) to Dog Lake ( 9240 ). Hud- 
sonian Zone; Subalpine Forest; talus slides; rock gardens. Lycaena cupreus. 
Donohue Pass: Inyo N F (11, 100). Arctic- Alpine; meadows, exposed rock. 
65. Elizabeth Lake: Cathedral Range (9500). Hudsonian Zone; Lodgepole Pine; 
meadows, willows, heather. Lycaena mariposa, Polites s. tecumseh. 

Gaylor Lakes: Above Tioga Pass Checking Sta. (10, 500). Arctic-Alpine; 
Alpine Fell- Fields, Includes Upper Gaylor Lake. Melitaea d. malcolmi. 
Glen Aulin: On Tuolumne River (7850). Canadian Zone; Jeffrey Pine; 
arid slope. 


2([); i -9€, 1963 


YOSEMITE BUTTERFLIES 


95 


68. Helen Lake: E of Kuna Crest (10, 896 ). Arctic-Alpine; White-Bark Pine. 

69 . Hoffmann, Mt. : W of May Lake (10,921). As above; flowers in gravel. 

70. Lambert Dome: N of Tuoltimne Meadows (9400). Hudsonian; forested. 

71 . Lyell Base Camps: Head of Lyell Fork (11,000), Arctic- Alpine; treeline, 

72. Lyell Fork Meadows: (9000). Hudsonian; Lodgepole Pine; willow stream. 

73. Ottoway Lake: Clark Range (8000). Canadian Zone; Red Fir Forest. 

74. Mammoth Peak: N end, Kuna Crest (12,225), Arctic- Alpine; exposed. 

75. Merced Lake Trail: From Merced Lake (7100) to Tuolumne Pass (10, 100) 
via Fletcher Creek, Canadian to Hudsonian; Coniferous Forest; Meadows. 

76 . Mono Pass: Inyo N F (10599), S of Tioga Pass, between Mt. Gibbs and 
Mt. Lewis. Arctic- Alpine; Alpine Fell-Fields, Hesperia miriamae. 

77. Pilot Peak: NW above Tioga Pass via Gaylor Lakes Trail (11, 100), Hud- 
sonian, Arctic- Alpine; Subalpine Forest, Alpine Fell- Fields, (See 81). 

78. Rafferty Creek: (8600- 10, 100). Hudsonian; Lodgepole Pine; Riparian. 

79 . Saddlebag Lake: Inyo N F (10, 050). Upper Hudsonian and Arctic- Alpine; 
few trees, willows, grasses, some rock. Includes E above (10,500). 

80. Tioga Meadow: (9941). Montane Meadow E of Tioga Pass Checking Sta. 

Tioga Pass: (9941). Hudsonian Zone; Subalpine Forest; Glacial Moraine. 

81. Tioga Pass, N.W above: (10, 500- 11,000). Tree-line Association; rock 
gardens. Parnassius p. behrii, Pieris o. calyce, Oeneis c. ivallda . 

82. Tenaya Canyon: (4, 000-8,000, average 6,000), Canadian Zone; stream 
bed choked with firs, aspen, willows; exposed slopes (Philotes enoptes) . 

83. Thousand Island Lake: Inyo N F (9850), Hudsonian; Lodgepole Pine; sage. 

84. Tuolumne Meadows: On Tioga Road (8600). Hudsonian; Lodgepole Pine, 
extensive meadows. Site of Lembert Homestead; metropolis of C. behrii . 

85. Unicorn Peak: Cathedral Range (10, 849). Arctic- Alpine; boulder j\imble. 

86. Vogelsang; Trail from Vogelsang Camp (10, 100) to Vogelsang Pass (10,400) 
and below. Hudsonian, Arctic- Alpine; Subalpine Forest; Alpine Fell-Fields. 

East- slope localities visited during survey; mostly Mono Basin; all out- 
side Yosemite National Park: 

87. Bodie : 11 mi E of Highway 395 (8369). Pinyon; Great Basin Sage; Meadow. 

88. Bridgeport: On Highway 395 (6743). Willow Thicket; Riparian Association, 

89. Casa Diablo Hot Springs; Inyo N F, on Highway 395 (7198). Transition; 
Great Basin Sage Brush in Jeffrey Pine Forest; sparse Streamside Assoc. 
Conway Pass : On Highway 395 N of Mono Lake, 200 ft. below Summit 
(8136). Great Basin Scrub (Desert Peach, Chrysothamnus, Purshia); 
Montane Meadow; Aspen Grove. 

91 . Deadman Summit: Inyo N F, on Highway 395 (7734 or 8168). Transition; 
Jeffrey Pine Forest; Purshia, Artemesia, Chrysothamnus, Eriogonum. 

92 . Gull Lake: Inyo N F (7600), on June Lake circuit. Moist Montane Meadow; 
Willow Thicket; Brownie Thistle (Cirsium aca.ulescens^ A. n. apacheana. 

93 . June Lake: Inyo N F (7716). Junction of Highway 395 and June Lake Road. 
Transition; Jeffrey Pine Forest; Great Basin Sage, Neophasia menapia, 

Lee Vi ning Creek: Inyo N F (7000). Above turnoff to last campground on 
Tioga Pass Road. Dry Hillside; Manzanita; Ceanothus, Bitter Cherry. 

95 . Mono Lake- ; (6419). Many associations in close proximity: Freshwater 

Marsh, Salt Sink Margin, Willow Thicket, Aspen Grove, Moist Meadow. 
Mammoth: Inyo N F (7900). On Highway'395, about 2-3 mi below. Tran- 
sition; Jeffrey Pine Forest; Great Basin Sage Brush; Great Basin Scrub. 
L'u^dy^ Inyo N F (7766), Sage, Aspen, Willow. Collecting to 8700 feet. 
Warren Creek: Inyo N F (9000-9500). N fork of Lee Vining Creek above 
Tioga Road crossing. Canadian Zone; willowed stream, sage-aspen slope. 


96 


GARTH AND TILDEN 


/. Res. Lepid. 


THE LEFIJDOPTE^A FOUHDATUOH 

THE FOUNDATION exists as a charitable fund to provide per- 
manent security to the Journal, Funds taken in are placed in a place 
or places where they will obtain reasonable earning power together 
with safety. Regular membership in the Foundation is 10.00 per year 
(coinciding with the Journal volume); other memberships are avail- 
able. See the special form later in this issue. The Foundation will also 
maintain collections suitable for geographical or biological study; gifts 
are welcomed from anyone. No public exhibits are planned. The 
Foundation welcomes help in its cause; it does not feel competitive with 
any other group but rather complementary. Arrangements for pay- 
ment in foreign currencies are possible; write to the editor for in- 
formation. 


YOU ARE CORDIALLY INVITED 

TO BECOME A MEMBER OF THE LEPIDOPTERA FOUNDATION, WHICH IS 
DEDICATED TO THE ADVANCEMENT OF KNOWLEDGE OF MOTHS AND 
BUTTERFLIES THROUGHOUT THE WORLD, THROUGH THE PUBLICATION 
OF THE JOURNAL OF RESEARCH ON THE LEPIDOPTERA. 


Membership includes subscription to the JOURNAL and a donation of the tolance 
of the subscription to a fund which wiU be used for a permanent endowment. 


THE LEPIDOPTERA FOUNDATION 
1140 West Orange Grove Ave., Arcadia, California, U.S.A. 


Application for membership: 

Name.. . 


hereby applies for...................... 

membership in the Foundation. 


CLASSES OF 
MEMBERSHIP 


Regular 

$ 10 year 

Family 

15 year 

Contributing 

25 year 

Subscribing 

50 year 

Sponsor 

100 year 

Life^ 

250 for life 


®Life membership includes the Journal for your life; should any future change occur such 

that the Journal would cease publication, the full amount of the Life membership will 

be returned to you during your lifetime. 

The Foundation will receive bequests or gifts in the form of funds, property 
or coilertions. 


advertisement 


TAKE CHLOROCRESOL ON YOUR NEXT TRIP 


Since the publication of the Chlorocresol method, 
field collecting techniques have been changed radi- 
cally. The Chlorocresol method eliminates the need 
for time consuming field mounting and/or relaxing 
of specimens. With Chlorocresol, specimens are re- 
tained in a relaxed condition for an unlimited period, 
allowing mounting to be done at a more convenient 
time or place. Field mounted specimens which would 
normally occupy a standard insect box may be packed 
in a container only 5" x 5" x IV 2 ". With specimens 
in a relaxed state, damage is minimized in the event 


of rough handling. 

Catalog No. 182 

Chlorocresol, 50 grams $0.95 

Chlorocresol, 100 grams 1.75 

Chlorocresol, 200 grams 3.00 

Styrene box, 5" x 5" x per dozen 3.00 


Complete instructions are included with each 
shipment. 

Prices are F.O.B. Santa Monica, California. California 
residents please add 4 % sales tax. 

GENERAL ENTOMOLOGICAL SUPPLIES 



BIO METAL ASSOCIATES 

Box 61 

Santa Monica, California 


THE JOUI^NJAL OF KES1AP.CHJ 
©H THE LEPIJ©©FTERA 


YOSEMITE BUTTERFLIES: An ecological survey of the 
butterflies of the Yosemite sector of the Sierra Nevada, 
California John S. Garth and J. W. Tilden 











f 

f} 


■/ 


,/ 


•«. 




\ie 

f 

I 








% 


■ "■3"< 

' • . ^ V 


<. 




,4$‘: 


Volume 2 

Number 2 

September, 1963 





a quarterly published at 

1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
edited by: WILLIAM HOVANITZ 

THE PURPOSE OF THE JOURNAL is to combine in one source 
the worlc in this field for the aid of students of this group of insects 
in a way not at present available. THE JOURNAL will attempt to 
publish primarily only critical and complete papers of an analytical 
nature, though there will be a limited section devoted to shorter papers 
and notes. QUALITY WORK on any aspects of research on the 
Lepidoptera is invited. Analytical and well illustrated works are pre- 
ferred, with a minimum of long description. 

AUTHORS ARE REQUESTED to refer to the journal as an 
example of the form to be used in preparing their manuscripts. Illu- 
strations should be of the best quality black and white, or line draw- 
ings and should be pre-arranged by the author to fit a reduced size 
of 4” X 6 V 2 .'' Footnotes should be avoided; bibliography should be as 
indicated. Tables should be set-up for page size as indicated. Manuscripts 
in good form and requiring little work by the editor will naturally 
appear first. Authors, who wish drawings made for them, may submit 
rough sketches and will be billed for the cost, which will be very 
negligible. Work to be done on research grants should so specify. When 
possible, tabular matter should be typed on good paper with a carbon 
ribbon in a form suitable for a one-third reduction and in a size to 
fit 4” X 61/2." 

THE JOURNAL is not a suitable place for continued changes 
of nomenclature; unless the author is himself analytically studying a 
group from its biological point of view and finds a change necessary, 
the editor must ask authors to refrain from any changes from the 
McDunnough Check List unless superseded by a monograph published 
since that date. Popular books are not to be considered as giving scien- 
tific credence to any name. It is rare that name changes need be made 
and preference is given to old names unless in the editor s opinion 
sufficient evidence is given to warrant such change. 

SUBSCRIPTIONS should be sent to the above address. 

RATES are: $8.00 per volume, personal subscription (but see below) 
$12.00 per volume, institutional subscription. 

The personal subscription rate is included in the membership to the 
Lepidoptera Foundation indicated below. SPECIAL SERVICE TO 
FOREIGN ADDRESSES: THE JOURNAL will be mailed air mail 
or registered at cost to the subscriber, if so desired. 


Journal of Research on the Lepidoptera 


2 ( 2 ): 97425 , 1%3 


114© W. Orange Grme Ape., Arcadia, California, U.S.A> 
© C&pyright iftfj 


QUANTITATIVE ANALYSIS OF CERTAIN WING AND 
GENITALIA CHARACTERS OF PIERIS IN 
WESTERN NORTH AMERICA 

VINCENT C. S. CHANG' 

California Arhoretum Foundation, Inc., Arcadia, Calif., and 
Los Angeles StSe College, Los Angeles, Calif. 

All species of ^Pieris which occur in western North America were 
named in the nineteenth century at which time wing coloration and 
pattern were the most obvious characteristics used for identification. 
Great variations are present between various species of the genus Pieris 
with respect to these characters. Such variations have contributed to 
disagreement concerning the number of species which occur in North 
America. Some authors list seven species: P. rapae, P. napi, P. vir- 
girdensis, P. sisymhrii, P. beckeri, P. pmlodke and F. occidentdis ( Klots 
1932 , McDunnough 1939) . Abbott et ai ( I960) in a statistical study 
utilized six morphological characters which included the length and 
the width of the front wing, the length of the hind wing, body length 
and the degrees of melanization of the upper and lower sides of the 
wings of the specimens. They found that there is a continuous varia- 
tion in wing sizes, color and marking between P. protodice, P, occi- 
dentdis and P. sisymbrii. Such variations were found associated with 
geographical location and climate. They suggested that F. protodke, 
P, occidemtdis and F. sisymbrii were a single species. Klots (1961) in 
Hs ktest Pieris list, cited only six species, excluding F. occidentdis. 
Hovanitz (1962) combined the distribution of P. occidentdis • mA 
P, pfotodice in a geographical study because of uncertainty of how to 
separate them. Evidence for considering them as separate species is 
given in this paper. Hovanitz (1963) also indicated that F. mpi and 
P. vifgmiensis ought to be considered as a single species at the present 
time for nomenciatiiral purposes pending more complete evidence of 
their possible sympatric distribution. Since F. vwgimemis occurs only 
in the region east of the Rocky Mountains in North America, it is 
excluded from consideration in this paper. In the present paper, mainly 
morphological characters are studied, m^hich are intended to aid in 
the elucidation of the biological relationships between these various 
species of Pieris. 

^This work was supplementary to work aided by a grant from the National 
Science Foupdation. 


97 


98 


CHANG 


]. Kes. Lepid, 


MATERIALS AND METHODS 


Five hundred specimens have been studied from the collections 
gathered by William Hovanitz and the author throughout the region 
in recent years. Despite this fact, this study is not intended of an 
exhaustive geographical study. 

The wing characters used in this study include: (1) wing color 
and pattern^ of which the detailed description will not be given; how- 
ever, the most important characters are outlined in the text, and 
(2) wing venation, which is relatively uniform in the genus Pieris. 
Differences have been found in the length of the Ry, and in the location 
of R 2 in respect of M 2 and (3) the androconial scales which are the 
scent. scales found with the black scales in the dot on or near the discal 
vein on the male fore wings. 

The characters used for the male genitalia include ( 1 ) the uncus, 
(2) the juxta, (3) the saccus and (4) the penis. For the fem?le 
genitalia, only the signum bursa has been used; this is a chitinized 
structure on the corpus bursa. 

The overall size of the insects influences the size of their internal 
structures. To make this size difference of negligible influence, the 
simple ratio: the length of the left front wing against the length of a 
particular structure has been used for aiding in the elimination of 
such possible differences. 


Ratio 


the length of a given structure ^ ^00 
the length of left front wing 


The terminology of structural parts employed here has been adapted 
from Klofs (1931). The color terms whenever used in the text follow 
the "Dictionary of Color” by Maerz and Paul (1930). 

L P. rapae and P. napi 

Color and pattern (fig. la): P. rapae and P.-napi are different 
from other species by lacking a dot on the discal vein of the fore wings. 
Most samples of P. rapae and P. napi are easily separated, except the 
all-white forms of both species. 

Venation: The mean ratio (table 1) between the length of the 
left fore wing and the length of the discal cell are statistically differ- 
ent for these two species. The probability that the difference might 
be due to chance alone is less than 0.001 when the degree of freedom 
is 45 ( as tested by ) . The histogram ( fig. 3 ) shows however that 
the range of ratios is continuous from one species to another. In 
actual fact, however, 00 overlap has been observed in the investigation 
between these two species though it may occur when the sample size 
is larger. 

Location of the discal cell, and method of measuring venation are 
illustrated in fig. 2. 


2(2):97-l25, 196} 


MORPHOLOGY OF PIERIiS 


99 



Fig. la. 


Pieris rapae and Pieris napi. Male on left, female on right. Upper 
four are upper side;lower four are lower side. 


100 


CHANG 


]. Res. Lepid. 



Fig. lb. Pieris beckeri and Pieris sisymhrii. Male on left, female on right. 
Upper four are upper side, lower four are lower side. 


2(2):97.12i, 1963 


MORPHOLOGY OF PIERIS 


101 



Fig. Ic. Pieris protodice and Pieris occidentalis. Male on left, female on right. 
Upper four are upper side, lower four are lower side. 


102 


CHANG 


]. Res. Lepid. 



Se D, = Standard deviation; S, E. = Standard error 


2 ( 2 ): 97 - 125 , 1963 


MORPHOLOGY OF PIERIS 


103 


Androconia (fig. 4, 5, table 2): Both P. rapae and P. napi have 
androconial scales with broadened tops and two lobes. A difference is 
found on the width of the scale’s stalk which for P. rapae is narrower 
than P. napi. The probability as shown by the 'P” test that the differ- 
ence between the mean ratio of these two species is due to chance 
alone is less than 0.001 for a degree of freedom of 40. The histogram 
(fig, 5) shows that there is a wide gap between the ratios. This is one 
of the characters that can be used to separate the all white forms of a 
P, rapae and P. napi. 

Male genitalia: (1) the uncus (figs. 6, 7, table 3) is more slender 
and less curved at the ventral side for P. rapae and P. napi than for the 
other four species in the genus Pieris. The ratios: the length of the 
uncus against the length of the fore wing range from 2.07 to 2.84 for 
P. rapae; and frorn 2.69 to 3.05 for P. napi. The probability that 
this difference might be due to chance alone is 0.025 for a degrees 
of freedom of 18 as tested by t: 





Fig. 2. Location of discal cell and points of measurement 


DISCAL CELL 


104 


CHANG 


I. Ii,s. LcpiJ. 





j 


O 

Q 

(a 

kj 


k 

Ui 

>J 



1 

a: 

-o 

g 




lo 

(%J NOUVindOd 



Pier is eapf-' 



'Pier is sisymbrfi 



Fig, 4. Androconial scales (400x) 


ANDROCONIA 


106 


CHANG 


]. Res. Lepid. 



(%) NOlivnndOd 


o 

o 

X 





istogram illustrating the differences in ratios distribution between 
, rapae and P. napi 


107 


2(2):97.12K 196} MORPHOLOGY OF PIERIS 


m 

d 



















•3 


4J 












m 



m 


+ 





04 



o 



d 


Rj 


O 



o 


04 

o 


■,Q 


• 



• 





O 


o 



o 


O* 

m 


u 








O 



















00 

•M 




in 



m 


CO 

rt 


+i 


• 

04 



• 

eo 


1—4 











nd 










d 










nt 


W 

if» 



t^ 

I— 1 



m 




O 

o 

f-i 

O 

o 

9) 

u 



O 


• 

o 

• 

o 

o* 

o* 

o* 

n 










o 










d 


• 

00 


Nl 

cn 

cn 

04 

04 



Q 

1— 1 


r~i 

04 

cn 

04 

04 

cn 



• 


• 

• 

• 

• 

• 




o 


o 

O 

o 

O 

O 











(0 


o 








m 

os 

s ° 

1^ 


p—t 

1— 1 

m 

rsj 

o 

d 

o 


00 

r- 

04 

p— 4 


(0 

d 

u 

d 

® t; 

• 

ra 


04* 

• 

fM 

04* 

• 

fM 

• 

04 

o 

•fH 

!□ 

IS 
















? 

m 

O 


"S d 




g 

g 



,d 


"S) *5! 
d ^ 

0) 

rH 

d 2 

s* 

« 


g 

H 

d 

s 

g 

g 

-M 

bO 

d 

<u 

r—l 

d 


g 

sO 

• 

fM 

04 


g 

00 

• 

04 

C 

Tt* 

cn 

c 

o 

04 

d 

00 

t— < 

• 

04 

g 

m 

• 

04 

04 

g 

• 

cn 

04 

4) 

0) 


S -S 

















■M 










4) 












X 








00 

O 

"H 


4J 

bO 

1 


1 

g 

g 

1 

g 

d 

1 



r-l d 




w 


Q 

d 



d ^ 

O 

in 


O 

m 


00 

vO 




t^ 

m 

m 


m 



<y 

o* 


• 

o 

• 

o 

o* 

• 

o 

• 

o 

ro 










4) 

1^ 

rO 


o' 

o 


00 

11 

o 

37 

m 

04 

nJ 










H 









m 










•H 







•1-1 

d 


4) 

O 

rH 

n 



OS 

(U 

4) 

(li 



43 

c* 

•«H 

d 


d 

« 



o 


H-l 

i 

4) 

4< 

o 

•M 

Td 

tH 



<D 

Cb 


p. 

GO 

O 

o 

O 




Rj 


nS 

•H 

4) 

d 

U 






d 

00 

rQ 

P4 

O 


108 


CHANG 


J. Kes. Leptd. 


(2) The juxta (fig. 7) is quite different in shape between P. 
rapae and P. napi. The lower end of the juxta of P. rapae is triangular; 
however, in P. napi, this structure is reduced in size and not hollowed 

out. 

(3) The saccus and penis (fig 8, table 4) show no significant 
differences between P. rapae and P. napi, 

Signum bursa ( fig. 8 ) : This structure is again a good character 
to separate P, rapae and P, napi. The signum bursa of P. rapae is like 
a pair of kidneys; the central part is less toothed. In P, napi, it has 
a long unchitinized tail which is so distinctive that there is no problem 
in separating this species from others in the genus Pieris. 

The samples of P. rapae came from ( 1 ) Arcadia, Los Angeles Coun- 
ty, California (5 individuals), (2) Newport Beach, Orange Co., Calif. 
(5), (3) Riverside, Riverside Co. Calif. (5), (4) Bishop, Inyo Co. 
Calif. (5), (3) Klamath Falls, Klamath Co. Oregon ( 5 ), and (6) 
Satus Creek, Yakima Co. Washington ( 5 ) . The samples of P. napi 
came from (1) Lopez Canyon, San Luis Obispo Co. Calif. (5), (2) 
Berkeley, Calif. (1), (3) Utah (1), (4) Flurricane Ridge, Olympic 
National Park, Wash. (10), (6) British Columbia (2), (7) Yukon 
Territory (6), and (8) Alaska (3). 



Fig. 6. The lateral view of male genitalia 


2(2):97-i25, 196) MORPHOLOGY OF PIERIS 




Fig. 7. Male genital structures 


109 


Table 4 Ratios between length of various genital structures and length of front wing 


110 


CHANG 


/. Res. Lepiil. 


Probability 

+ 

o 

+ 

« 

o 

o 

o 

o* 

■4^ 

00 

cn 

• 

O'- 

« 

o 

fNl 

« 

(M 

W 

m 

0, 09 

0. 07 

60 *0 

01 *0 

0* 04 

0. 04 

Q 

m 

fM ^ 

• • 

o o 

0. 28 

0, 24 

0. 31 

0. 19 

Mean of 

ratio 

rj o 

• m 

O t-” 

« • 

00 m 

» « 

ra fP) 

Mean length 
of front wing 

g 

g 

O' 

m 00 

V 

tM 

00 

r- 

O* ro* 

22.96 

23. 54 

Mean length 
of saccus 

0, 51 mm 

0. 51 

« — ) 

• • 

O O 

00 

• # 

o o 

o 

a- r- 

00 00 

53 

30 

Species 

rapae 

napi 

sisymbrii 

beckeri 

protodice 

occidentalis 


a 

ro 

O 


Probability 

O 

o 

o* 

o 

o 

• 

o 


cn 




00 


• 

# 

4^ 

r«- 

r- 




W 


fM 


PO r~4 


m 

# « 

• » 

m 

o o 

o o 




d 

o o 

o o 


—1 Ifi 


m 

* « 

# « 

m 

^ o 

pH 

o 




r-t 

pH s-H 


m * 

9 # 










bJO 



d 

g g 

s g 

0 

M H 

f=^ 

M N 

f=s H 

fH 

6 g 

g g 

a 

a- 

1^ 

d 

0 fO 


00 fM 


o* o* 

O* O* 

rd 



4^ 



m bo 

fl d 



R G 

s s 

0 ^ 

6 B 

G 6 


N N 

G K 


PO Pj 

fO O' 


r- 00 

r~ 

0 o 


« ■ 

^ ij 

o ^ 

O fM 


rM ra 

fM fM 




o* 

f%J p>4 

<M 


P-t .-1 

pM fO 



^r4 



0 



N y 


■§ S 

■d ^ 

M 

;• ■:: 

g o 

0 

ffl o 

m P 

•rt 

^ 0 


u 

rts 

M ^ 

m ^ 

M 

0.* 

d d 


2(2)^97-12K ^963 


MORPHOLOGY OF PIERIS 


111 




P E N i S 


S i 6 MU M 
b’ U R S A 



pier is sisymtrii 




iic e.. 



Fig. 8. Male and female genital structures 


112 


CHANG 


J. Res. LcpiJ. 


II. P. sisymbrii and P. beckeri 

1. Color and pattern (fig. lb); These two species are easily 
separated by the color and the pattern on the wings. P. beckeri has 
the intensified moss green color and P. sisymbrii has the biskra data 
color along the vein on the underside of the wings. 

2. Venation (fig. 9, table 5): The length of between P. 
sisymbrii and P. beckeri differs. The ratio of fore wing length over 
R 3 for P. sisymbrii ranges from 2.34 to 5.88; for P. beckeri ranges 
from 0.95 to 2 . 62 . The probability that the mean for the two popu- 
lations could have been drawn by chance from the same population 
is less than 0.001 when degrees of freedom is 24 as tested by t. 

3 . Androconia (fig. 4): The scent scales of P, sisymbrii and 
P, beckeri are similar but unlike P. rapae and P. napi. The tiny, lobe- 
less scales of P, sisymbrii are nearly parallel at the lateral border, and 
the roots are limited to the base. The scales of P. beckeri are usually 
irregular shaped and the roots are often occupied half the length of 
the scale. 

4. Uncus (fig. 7, table 3) ; The uncus of P. beckeri and P. nsymbrii 
have hairy processes at the ventral side, particularly in P. sisymbrii. 
The actual length of the uncus in the two is much the same, but the 
ratio is different (probability 0.005; d.f. 20) due to the constant 
smaller size of wing length in P, sisymbrii. 

5 . Juxta (fig. 7): The juxta of P. beckeri and P. sisymbrii are 
similar except the apex is slightly curved upward in P. beckeri. 

6. Saccus (fig. 8, table 4): There is little difference in shape or 
size with respect to this structure. 

7. Penis ( fig. 8 ) : The shape of the penis in P. sisymbrii and 
P. beckeri is like the penis of P. rapae and P. napi. However, it is 
distinctly different from the penis of P. protodice and P. occidentalis. 
The basal protuberance of the penis of P, beckeri and P. sisymbrii is 
insignificant. But it rises abruptly for P. protodice and P. occidentalis. 

8. Signum bursa ( fig. 8, table 6 ) : The signum bursa of P. sisym- 
brii and P. beckeri is stick-shaped, with large, heavily chitinized teeth. 
The difference between the ratios is probably caused by chance error 
due to the extremely small sample of P, sisymbrii examined, rather than 
to a real difference. 

The samples of P. sisymbrii came from ( 1 ) Kelso Valley, Kern 
Co. Calif. (2), (2) Roads End, Tulare Co. Calif. (1), (3) Kings 
Canyon, Fresno Co. Calif. (2), (4) Hurricane Ridge, Olympic Na- 
tional Park, Wash. (6), and (3) Whitehorse, Yukon Territory (1). 
The samples of P. beckeri came from ( 1 ) Long Valley, Mono Co. 
Calif. (1), (2) Mono Lake, Mono Co. Calif. (2), (3) Gardnerville, 
Douglas Co. Nevada (2), (4) Minden, Douglas Co. Nevada (2), 
(3) Doyle, Lassen Co., Calif. (2), and (6) Sams Creek, Yakima Co. 
Wash. (3). 


Table 6 Ratios between length of various genital structures and length of front wing 


2(2):97-I2'>, l')6) 


MORPHOLOGY OF PIERIS 


IH 


d 

bO 

*r4 

m 


Probability 

0 

• 

0 

0 

0 

• 

0 


sO 



00 

• 


• 



CO 

I— < 

» 

ra r- 


N 

CO irt 

CO m 


0 0 

0 0 

• 

• • 

• • 

CO 

0 0 

0 0 



m 

Q 

m 

0 m 


0 

■-I (S3 

• 

• • 


CO 

0 0 • 

0 0 

0 






4J 






Pi 



Vi 



0 



d 

00 ^ 

00 CO 

RS 

OJ 00 

fH 0 

0) 

• • 

• * 


CO <M 

CO 

bO 




P 



N 

g 



r^ 

(S3 


00 00 

CO 0 

§ 5 

• • 

fsl ^ 

• • 

CO 


(S3 (S3 

(S3 (S3 

l-s 



Rl 



at 

Pi 



^ J 

4J 

g 


be 

d 


d g 

g 


0 p 

r-< ^ 

m so 

^ vO 

d 

t- 


5 -fI 

• • 

• • 

(e an 

d) 

0 0 

0 0 

Vi 

0 






d 


10 m 

Z 

(S3 sO 

CO (S3 



00 





*r4 

t—i 

<e cj 

m 

0) 

^ ^r4 

■S 

nd 0) 

*y 

i ^ 

0 

4-> -fI 

4) 

m 0 

0 t» 

P. 

* 9) 

P( 0 

CO 

m ^ 

p 0 


Q 

rd 

m 

PI 

0) 

■U 

d 

O 

d 

rt 

u 

o 


W 


CO 


CO 


pcj 


•rH 'd 


d p 


o 

d) 

P4 

CO 


0 

o 

o 

o 

Pi 

p. 




114 


CHANG 


]. Res. Lepicl. 


III. P. protodice and P. occidentalis 

1. Color and pattern (fig. Ic, table 7): The color and pattern 
of these two species are many times undistinguishable, especially in 
comparison between the high temperature form of P. occidentalis and 
the low temperature form of P. protodice. The primary difference in 
the pattern, besides the gross one of more extensive black pigment on 
P. occidentalis males than on P. protodice males, is in the size and 
shape of the dot in the tornus area near the inner margin on the 
underside of the fore wings. The dot on P. protodice is large, square 
shaped and not clearly outlined, part of the dot being sometimes sub- 
merged on the uppersides. The dot of P, occidentalis is small, relatively 
clearly outlined, usually not submerged. The ratios for the width of 
the tornus dot range from 8.73 to 15.38 for P. protodice; and from 3-95 
to 8.51 for P. occidentalis. The probability that this difference is due 
to chance alone is less than 0.001 for the degrees of freedom of 60 
by the t test. 

2. Venation (fig 9, 10, table 5); The R:^ veins of P protodice 
and P, occidentalis are quite short, sometimes being completely miss- 
ing. The mean ratios of wing length over length between the two 
species do not show much difference. (1.17 for P. protodice\ and 1.15 
for P. occidentalis). However, the difference is significant between 
P, protodice and P. sisymhrii in comparing the ratios despite ^the sim- 
ilarities in their wing color and pattern. The mean ratio for P. sismyhrii 
is 4.16; and for P. protodice it is 1.17. That the difference as indicated 
might be due to chance is less than 1/1000 for the degrees of freedom 
of 46 by the t test. 

In P, protodice, the intersection of Rl>, Rs+Mi (point "A”) is usual- 
ly above point "B” where M 2 joins that vein; hov/ever, point "A” is 
usually on or below point "B” in P. occidentalis. The distance between 
point "A” and point "B” is measured and plotted on the diagram 
(fig. 11) against the ratio obtained from the width of the tornus dot 
to the length of the fore wing. When point "A” is above point "B” 
the measurement is given a "-b” sign. When point "A” is below point 
”B” the measurement is given a '' — ” sign. The diagram (fig. 11) 
shows that P. protodice and P, occidentalis are well separated by these 
two characters. Few exceptional dots on the diagram as the measure- 
ments between point "A” and point "B” having a ” — ” sign for P. 
protodice or vice versa for P. occidentalis are from the specimens col- 
lected at the overlap zone of these two species. 

The width of tornus dot and the location of R 2 in respect of 
M 2 are two pirmary characters used in the investigation to separate the 
two species. 

3 . Androconia: Thirty-five specimens of male P. protodice and 
twenty-eight specimens of male P. occidentalis from ten different 
localities were checked, but no androconial scales have ever been found. 
Similar results were also observed by Bernard! ( 1947) . 


2 ( 2 ); 97 - 125 , 1963 


MORPHOLOGY OF PIERIS 


115 





P . protodice 



Fig, 9. Venations showing the differences in the length of R 3 


116 


CHANG 


]. Res. Lepid. 



prim ary 
P. protodice 




Fig. 10. Venations showing the left fore wing and the differences in the 
location of point "A” in respect to point "B” between P. protodice 
and F. Occident alts 


2(2):97-l2^, 196} 


MORPHOLOGY OF FIERI S 


117 


4. Male genitalia: (1) Uncus and juxta (fig. 7, table 3 ): No 
difference is found between P, protodice and P, occidentalis in respect 
to these two characters. 

(2) Saccus (fig. 8, 12, table 4): The length of saccus is statis- 
tically different between P. protodice and P. occidentalis. The mean 
ratio (length of fore wing over length of saccus) for P. protodice is 
2.68; for P. occidentalis it is 3.45. The probability is less than 0.001 
that the difference may be due to chance when the degrees of freedom 
is 83 as shown by the t test. 

The .scatter diagram (fig. 12), which shows each individual ratio 
of these two species against its own wing length, shows that overlap 
is present in this character despite the difference in mean ratio. This 
may be due to the ‘variation in wing size caused by geographical and 
climatical difference. In cooler areas, the butterfly is usually smaller 
and in warmer areas, larger. If some specimens of P. occidentalis are 
from the extreme southern locations and some specimens of P. protodice 
from the extreme north, the gap in ratio between these two populations 
is reduced, and overlap occurs. This point is demonstrated by fig. 14. 
If the specimens of P. protodice and P, occidentalis are from the same 
locality and collected at the same time, these two species are usually 
separable. 

( 3 ) Penis ( fig. 8 ) : The penis of P. protodice and P, occidentalis 
is strongly curved at the basal part and this difference makes these two 
species easily distinguishable from other species in the genus Pieris. 
The penis size in P. protodice is smaller than in P. occidentalis. 

5. Signum bursa (fig. 8, table 6): The signum bursa of P, pro- 
todice and P, occidentalis is similar to P, sisymbrii, but with smaller 
teeth and less chitinization. The length of signum bursa in P. protodice 
is shorter than the one in P, occidentalis': The mean ratio (length of 
fore wdng over length of signum bursa) for P. protodice is 3.18 and 
for P. occidentalis is 4.03. The probability that this difference is due 
to chance alone is less than 0.001 when the degrees of freedom is 60 
by the t test. The scatter diagram (fig. 13) for individual ratios shows 
that there is no overlap present between these two species with respect 
to this character. 

The mean ratio of each population at a given locality for P. proto- 
dice and P. occidentalis is also plotted on a diagram (fig. 14) which 
shows that the ratios of the specimens from northern localities such 
as Alaska are larger than the ratios of the specimens from southern 
localities. The differences in ratios for the specimens from various 
southern localities are not significant. This is porbably due to the 
closeness in distance between the localities and to similarities in en- 
vironment. This diagram illustrates however that the ratio is not every- 
where unchanged. The ratio gradually increases in the more northern 


VENATION 
P. PROTODICE (k) 

P. OCCIDENTALIS { •) 


118 


CHANG 


/. Res. Lcpid. 



001 X 


iOO SnNMOi HiOIM 


Fig. 11. The measurements between point "A” and point "B” (see fig. 10) 
plotted against the ratios obtained from the width of the tornus dot 
to the length of the fore wing 


S AC C U S 


2(2):97-125, 196} 


MORPHOLOGY OF PIERIS 


119 


»(+++ 


-++ + + + 


44 


4- 4f 4-4- 




Q. (x: 


MJ JO H 19 N 3 1 


00! X 


snooir s jo h i 9N3 i 


Fig. 12. Scatter diagram showing the individual ratio of saccus length to the 
length of fore wing plotted against the individual saccus length in 
P. protodice and P. occidentalis 


V s y ns 


120 


CHANG 


/. Ke%. Lepid. 


o 



Q 

^ k 


k, 

Q 

SS 

!>> 

© 


<0 



o 


to 



o 


© 


00 ! X ^ ^ NI9N3 1 


8 § J O HI 9N31 


Fig. 13. Scatter diagram showing the individual ratio of the length of signum 
bursa to the length of fore wing plotted against the individual 
length of signum bursa in P. protodice and P. occidentalis 


2 ( 2 ) -. 97 - 125 , 1963 


MORPHOLOGY OF PIERIS 12 i 



■M'J HI3 1 N¥3m 


MIS M3 7 NV3M 


MEAM LENGTH OF SIGNUM BURSA (mm) 

Fig. 14 Scatter diagram showing the mean ratio of the length of signum 
bursa to the length of fore wing in each- locality plotted against the 
mean length of signum bursa 


122 


CHANG 


J. Res. Lepid. 


localities. The ratios of two species may overlap at different localities 
though they do not do so at the same location. 

The samples of P. protodice came from ( I ) Laguna Canyon, Orange 
Co. Calif. (11), (2) Moorpark, Ventura Co. Calif. (4), (3) Inde- 
pendence, Inyo Co. Calif. (2), (4) Big Pine, Inyo Co. Calif. (2), 

( 3 ) Mammoth Lake, Mono Co. Calif. ( 1 ) , (6) Mono Lake, Mono Co. 
Calif. ( 4 ) , (7) Coleville, Mono Co. Calif. (5) , (8) Gardnerville 
Douglas Co. Nevada (1), (9) Minden, Douglas Co. Nevada (1), 
and (10) Doyle, Lassen Co. Calif. (4). The samples of P. occidentalis 
came from (1) Long Valley, Mono Co. Calif. (1), (2) Mammoth 
Creek, Mono Co. Calif. ( 2 ) , ( 3 ) Mono Lake, Mono Co. Calif. ( 2 ) , 

(4) Coleville, Mono Co. Calif. (1), (3) Gardnerville, Douglas Co. 
Nevada (11), ( 6 ) Doyle, Lassen Co. Calif. ( 3 ) , ( 7 ) Klamath Falls, 
Klamath Co. Oregon (4), (8) Satus Creek, Yakima Co. Washington 
( 2 ) , ( 9 ) British Columbia ( 1 ) , and (10) Alaska ( 1 ) . 


GEOGRAPHICAL DISTRIBUTION OF P. PROTODICE 
AND P. OCCIDENTALIS 

The geographical distribution of all species in the genus Pieris in 
North America except P. protodice and P, occidentalis is thoroughly in- 
vestigated by Hovanitz (1962) (fig. 15). The data here presented are 
mostly gained by extensive search throughout the range of P. protodice 
and P. occidentalis in West Coast conducted by William Hovanitz in the 
summer of 1962. P, protodice is most abundant in the south, from 
the southern tip of Baja California northwards, gradually reducing its 
population and being replaced by P. occidentalis. The northern border 
of P. protodice is still unclear. At present this species is only in Cali- 
fornia and Nevada southwards. There is no record in Oregon and north- 
ward. P, occidentalis extends from central Alaska along the Rocky 
Mountains and the Cascade Mountains southwards into the Sierra 
Nevada Mountains (and the Rocky Mountains). The most southern 
locality known from our data at present in the Pacific coast area is 
Long Valley, Mono Co., Calif. It is very likely this species may be 
found at more southern places on high Sierra Nevada Mountains. 

P, occidentalis and P. protodice overlap from Long Valley, Mono 
Co. Calif, northward to Doyle, Lassen Co., Calif. Further investiga- 
tion may extend the overlap zone more northward even to Washington. 

The habitats of these two species are very similar; both like sunny, 
open, grassland. P, protodice prefers warmer temperatures and semi- 
desert conditions. It feeds on Brassica nigra^ Caulanthus sp., Lepidium 
densiflorum, Sisymbrium altissimum, Cleome lutea, and T hely podium 
lasiophyllum. P. occidentalis prefers relatively cool temperatures. It 
is usally found at places relatively moist, as near a lake, ditch or farm 
land. The food plants of P. occidentalis are nearly the same as for 


2(2):97-l2S, 196} 


MORPHOLOGY OF PIERIS 



Fig. 15. 


Map showing the distribution of P. protodice and P. occidentalis 
western North America 


124 


CHANG 


). Res. Lct)iJ. 


P. protodice except that Cleome Intea has not yet been recorded. 

P. protodice and P. occidentalis are found flying together in the 
overlap zone. The males of these two species at a particular locality 
are usually separable by the marking and color on the wings, in which 
P. protodice is less darkly marked than P. occidentalis. 

DISCUSSION AND CONCLUSIONS 

The present data indicates some clear structural differences be- 
tween the six species of Pieris. P. rapae and P. napi are similar in 
many characters, such as uncus, saccus, and the color and pattern on 
the wings, but they also can be distinguished by their differences in 
pattern and the signum bursa, juxta and androconial scales. P. sisymbrii 
and P. heckeri are similar in their uncus, saccus, androconial scales 
and signum bursa, but can usually be distinguished by wing pattern 
and venation. P. protodice and P. occidentalis can be distinguished by 
wing pattern, wing venation, structural differences in the length of 
saccus and signum bursa only in a statistical study. These two species 
also differ in their geographical distribution pattern, but overlap greatly 
in a sympatric area. 

P. sisymbrii differs in many characters from P. protodice, such as 
wing venation, genital structures and others. The most significant 
difference is the presence of androconial scales, v/hich are absent in 
P. protodice and P. occidentalis. 

The results serve to show that P. protodice and P. occidentalis are 
sympatric species, an uncertainty which led Hovanitz (1962) to com- 
bine their geographical distributions into one map. The other species 
of the genus Pieris in Western North America are P. rapae, P. napi. 
P. sisymbrii, and P. beckeri. It is always possible that even these may 
be found at some time in the future to be subdivided into cryptic 
species by some characters not yet discovered. 

ACKNOWLEDGEMENT 

The author wishes to express his deep appreciation to Dr. William 
Hovanitz and Dr. E. 1. Schlinger for their help and constant advice 
during the entire study. 


2(2):97-l2‘>, 196} 


MORPHOLOGY OF PIERiS 


125 


LITERATURE CIlTD 

ASAKAWA, F. 1961. Morphological observation on the androconia in the 
butterflies. (Ja) New Ent. 10(2): 7-15. 

ABBOTT, W., L. S. DILLON, and R. R. SHRODE. I960. Geographic 
variation in Pieris protodice Boisduval and Leconte ( Lepidoptera : Pieri- 
dae). Wasmann J. Biology ^ 18: 103-127. 

BERNARDI, G. 1947. Revision de la classification des especes holarctiques 
des genus Pieris Schr. et Pontia Fabr. (Pep., Pieridae). Miscellanea En- 
tomologica, Paris 44: 65-80. 

HOVANITZ, W. 1962. The distribution of the species of the genus Pieris 
in North America. /, Res. Lepid. 1 (1): 78-83. 

• . 1963. The relation of Pieris virginiensis Edw. to Pieris napi 

L. Species formation in Pieris.^ J. Res. Lepid. 1 (2): 124-134. 

JACKSON, R. A. 1946. Causes for seasonal variation in the number of 
Lepidoptera. Proc. & Trans. S. London Ent. & Nat. Hist. Soc. 1945-46: 
43-51. 

KLOTS, A. B. 1931. A generic revision of the Pieridae (Lepidoptera) to- 
gether with a study of the male genitalia. Entomologica Americama 7 
(new series) (3): 139-204; (4): 205-243. 

KLOTS, A. B. 1961. Family Pieridae. In "How to know the butterflies” 
edited by Ehrlich, P. R. and A. H. Ehrlich. Wm. C. Brown Co. Du- 
. buque, Iowa. 

MAERZ, A. and M. R. PAUL. 1930. A dictionary of color. 1st Ed. Mc- 
Graw-Hill Book Co. New York. 

MARIANI, M. 1937. Anatomia e fisiologia degli organ! genitali femminili 
delle Pieris Schrk. (Lepidoptera, Pieridae). Festschrift fur Prof. Dr. 
Embrik Strand. 3: 434-450. 

McDUNNOUGH, J. 1939- Check list of the Lepidoptera of Canada and 
the United States of America. Mem. So. Calif. Acad. Sci. 11(1) : 1-171. 

McHENRY, P. 1962. The generic, specific and lower category names of 
the Nearctic Butterflies. Pat 1. The genus Pieris. J. Res. Lepid 1(1): 
63-71. 

OKANO, M. 1951. Comparative morphology of the male genitalia of 
Japanese Pieridae. Ann. Report Gakougei Eac. livate Univ. 2: 38-46. 

PETERSON, B. 1963. The genitalia of some colias species. /. Res. Lepid. 
1 (2) : 135-156. 

QUERCI, O. 1936. Notes on Pontia protodice Boisduval and Leconte (Lep: 
Pieridae). Trans. Amer. Ent. Soc. 62: 37-47. 

TUXEN, S. L. (editor). 1956. Taxonomist’s glossary of genitalia in insects. 
Ejnar Munksgaard, Copenhagen. 

WARREN, B. C. S. 1961. The androconial scales and their bearing on the 
question of speciation in the genus Pieris, (Lepidoptera.) Ent. Tidskr. 
82(3/4) : 121T48. 


126 


CKANG 


). Res. Lepid. 


BIOGRAPHICAL SKETCHES 


CHANG, VINCENT CHUEN SUN 

[2855 Nina St, , No, 2, Pasadena, California] 

Born: Shanghai, China, March 17, 1935 

B„ A, : The Agriciiltural Institute of Nanking, Nanking, China, 
1957 


Graduate student: University of California, Riverside and Los 
Angeles State College 

Position: Research Assistant, California Arboretum Foundation, 


Inc,, 1959-63 

Interests: All aspects of Pieris, 
including ecology, phy- 
siology, and morphology 
Photography of activities of 
butterflies 

Publications: A series of papers on 



food preferences of Pieris 


Journal of Research on the Lepidoptera 


2 ( 2 ) : 127 - 136 , 1963 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
© Copyright 196 } 


GENETIC AND ENVIRONMENTAL VARIATION 
IN PIERIS BRASSICAE 

BRIAN O. C. GARDINER 

Agricultural Research Council, Unit of Insect Physiology, 

34A, Storey’s Way, Cambridge, England 


INTRODUCTION 

The principal form of variation in Pieris hrassicae L. consists 
of either a reduction or expansion in the amount of black present. In 
certain local populations the variation is constant and marked enough 
to be designated as a race, such as P. hrassicae cheiranthi in the Canary 
Islands. Marked variation in color, however, is exceedingly rare and, 
until now, only some half dozen specimens were known. 

In 1950 a stock of P. hrassicae originating from Cambridge was 
established at the Unit of Insect Physiology and has been continuously 
bred ever since. This stock has already been reported on (David & 
Gardiner 1951; David 1957). In 1959 another stock, originating from 
Oxford, was established as a separate culture and was also continuously 
brooded under the same conditions as the Cambridge stock. 

Until i 960 the amount of variation noticed in the Cambridge 
stock was simall, and confined to known variation in the black markings. 
This has already been briefly reported on (David «& Gardiner 1961). 

During the past three years, however, two color varieties have ap- 
peared, as well as the previously unknown albino form; known varieties 
also seem to have been more common, although this could well be 
due to more careful observation (fig. 1). 

The opportunity was taken to determine the genetics of these 
varieties and also to examine the combination of two pairs of allelo- 
morphs to produce a double recessive. It is believed that this is the first 
time such a cross has been done with a butterfly. 

From time to time specimens considerably smaller than the normal 
size turn up in various species of butterflies. In the case of P. hrassicae 
specimens of between 40-50 mmi. wing span are referable to ab. minor 
Ksienschopolsky. According to Frohawk (1934) normal wild caught 
specimens vary from 63 mm. in the male to 76 mim. in the female. 
In the continuously brooded culture kept at the Unit of Insect Physi- 
ology the size is 55-63 mm. in the male and 54-65 mm. in the female 
(David & Gardiner 1961). 


127 


128 


GARDINER 


/. Res. Lepid: 


Occasional specimens of ab. minor were noted in the Cambridge 
stock. At times entire batches have been on the small side. Specimens 
bred from these were normal. It occurred to the author that ab. minor 
might be an effect of starvation rather than a genetic effect. This 
has been investigated as will be described. 

DESCRIPTIVE 

Pieris hrassicae L. ab. coerulea ab. nov. 

The normal cream and green coloration on the underside of the 
wings is replaced by a pale blue color. On the upper side, the white 
areas of the wings have a pure white appearance and are rather thinly 
scaled; in normal specimens they are creamy white or off white in color. 

Larvae and pupae normal. The majority of the adults of this variety 
fail to eclose properly. Some specimens may emerge with striped 
scales, especially on the hind wing, which gives them a translucent' 
blue color, deeper in shade than normally formed coerulea. 

Holotype ^ , Allotype $ : Two specimens selected from the F i 
generation obtained by the pairing of two ab. coerulea which emerged 
from a continuous brooded culture kept in the Laboratory since 1950. 
In the author’s collection. 

Paratypes: Five pairs also selected from the Fi. Two pairs have 
been deposited in the Rothschild-Cockayne-Kettlewell collection at 
Tring, two pairs are in the author’s colletion, and one pair in the coL 
lection of Mr. H. Douglas Bessemer. 

Type locality: The original stock from which this aberration arose 
was collected in the vicinity of Cambridge, England, in 1950. 

Synonyms, ab pallida. Graham-Smith & Graham-Smith 
ab. anthrax. Graham -Smith & Graham-Smith 

Graham-Smith and Graham-Smith (19.30) after describing an- 
thrax state "Perhaps a specimen exhibited by Leeds (1909) with 
undersides of the hind wings a very distinct blue . . . are examples 
of this aberration.’’ 

Under pallida they state "A very marked example from Monk’s 
Wood, Hunts., is figured by Frohawk ( 1914, pi. 3, fig. 20) 

The specimen figured by Frohawk (1914) is stated by him to have 
been taken by Mr. H. A. Leeds in Monk’s Wood in 1906 and there 
can be little doubt that it was this specimen that was exhibited by 
Leeds at the Annual Exhibition of the South London Entomological 
and Natural History Society in 1909- 

Mr. A. L. Goodson has compared the present variety with the an- 
thrax and pallida in the R.-C.-K. collection at Tring and it in no way 
resembles either of them nor, in the author’s opinion, does it bear any 
resemblance to the illustrations of anthrax and pallida in the paper by 
Graham-Smith and Graham-Smith (1930). 


2(2):127-IH, 1963 


PIERIS BRASSICAE VARIATION 


129 


Mr. H. Douglas Bessemer, who has the original Leeds specimen in 
his collection, very kindly invited the author to visit him and examine 
it. There is no doubt that it is the same as the present variety and was 
wrongly' identified by Graham=Smith and Graham-Smith. 

Pieris brassicae L. ab. jmmi ab. nov. 

In May 1961 it was noticed that the underside of some butterflies 
being reared in the Laboratory was different in color to normal speci- 
mens, being a pale straw color compared to the normal greenish-yellow. 
The difference is more striking in freshly emerged living specimens 
than in dead set ones. The larvae are normal but the prepupae and 
pupae are a golden-yellow without any trace of the normal green. The 
pupae are also without the normal black speckling. Unlike some other 
color varieties the butterflies edose normally. 

Holotype , Allotype ’$ : Two specimens selected from the culture 
in May 1961. In the. author’s collection. 

Paratypes: Two pairs similarly selected. In the author’s collection. 

Type locality: The original stock from which this aberration arose 
was collected at Oxford, England, in August 1959. 

Pieris brassicae ab. alhinensis Gardiner 

This variety has already been described (Gardiner 1962). It is a 
simple recessive and is noted here since it was used with coemlea to 
obtain the double recessive. 

EXPERIMENTAL 

Most insects were bred according to methods already described 
(David & Gardiner 1952; David 1957) with the exception that when 
a single pair of butterflies was being paired a small nylon covered 
cage measuring 1 x 1 x lid feet was used instead of a large cage. A 
percentage of the larvae were also reared in 2 lb. jam jars at the rate 
of about 20 per jar. This enabled an accurate check to be kept on 
mortality. This was generally done with the F| broods, the larger 
numbers of the F 2 being reared in cages. No difficulties were encount- 
ered with either pairing or oviposition nor, apart from the Fi of ab. 
jauni, was there any appreciable mortality of the larvae. Due to limit- 
ations of time, space and food, it was only possible to rear a small 
percentage of the total ova obtained. Since a single fertile brassicae 
was found to lay over 500 ova, the potential F 2 was some 125,000 in- 
dividuals, an impossible number to rear. 

ab. coerulea 

In November 1962 a female emerged in the Cambridge stock. She 
was paired to a normal male from the same batch and the resulting 
ova were reared to give an Fi of normal insects. These were crossed 
and a selection of the resultant ova reared. 


130 


GARDINER 


]. Kcs. Lepiil. 


The F 2 produced 347 normal and II 6 coerulea butterflies, a ratio 
of 2.99 : 1, a very good agreement with the expected 3 : 1 ratio for 
a simple recessive. The majority of the coerulea however failed to 
eclose normally and remained stuck by the wing tips to the pupal case. 

While this brood was being reared, further coerulea emerged from 
a batch of the stock Cambridge butterflies. Two pairings v/ere obtained 
and all the Fi and subsequent generations were coerulea. This aberration 
is therefore a recessive. It’s extreme rarity under natural conditions 
can be explained by the difficulty the butterflies have in getting free 
of the pupal case. 

The blue color is due to an absence of the normal yellow pigments 
and is present normally in the wing-membrane. This can be shown 
faintly by rubbing off the scales of a normal hrassicae between finger 
and thumb. A very similar appearance to coerulea can also be pro- 
duced by immersing a normal hrassicae in the bleaching agent sold, 
under the name of 'Parazone.’ The question of the chemistry of this 
and other color forms of hrassicae is at present under investigation 
by a colleague of the present author and will be dealt with in a later 
paper. 

The failure to eclose is more marked in the males than the females. 
In the first two generations only about two per cent of the specimens 
could be described as immaculate. About 50 per cent fail to clear the 
pupal case and are stuck by one or both fore wing tips; others get 
clear but one or more wings fail to expand. In many instances there 
is extensive stripping of the scales, especially of the hind wing, which 
then has a deep blue translucent appearance. 

After several generations the percentage of normal eclosions rises 
to about 80 per cent. This is similar to the trend that occurred in 
alhinensis (Gardiner 1962). Since 1961 alhinensis has been passed 
through a further ten or twelve generations and the percentage failing 
to eclose normally is now negligible, and there seems little doubt that 
coeridea will eventually reach this stage. 

While failure to eclose is a semi-lethal factor linked, initially, 
with coerulea it is also partially controlled by the temperature at which 
eclosion takes place. Two experiments were done on this, using the Fo 
generation of the pure line in the first and alhinensis in the second. 

A cage of coerulea pupae was kept at 20 °C. until 12 normal 
(5.7%) and 198 crippled specimens had emerged. The remaining 
pupae were than transferred to 12.5°C. and at this temperature 38 
normal (27.0%) and 103 crippled specimens emerged. 

In the second experiment alhinensis was used, half the pupae being 
at 25°C. and half at 12.5°. At 25°C. 15 normal (30.0%) and 35 
crippled emerged while at 12.5°C. 28 normal (58.4%) and 20 crip- 
pled emerged. 

These results clearly show that failure to eclose is more marked 
at higher temperatures. 


2(2):127-1U, / 96 .! 


PIERIS BRASSICAE VARIATION 


131 


ab jaunt 

These were noted in May 1961 in a stock obtained from Oxford 
which had been bred for a number of generations in the Laboratory. 

A pair of these butterflies was mated and 100 of the ova reared. 
In this generation the mortality, mainly granulosis virus, was 76%. 
The 14 adults obtained were all of the variety. These were paired at 
random and 100 Fo ova reared to produce 97 butterflies, also of the 
variety. 

After the Fi males had paired with their sisters they were paired 
with normal females and a percentage of the ova laid were reared to 
produce all normal butterflies. These were mated at random to produce 
the F:> of which a percentage of the total ova laid were reared, with 
little mortality. This generation produced 100 normal and 34 ab jauni 
butterflies. This is a 3 : 1 ratio. 

This variety seems to be due to the lack of the normal greenish 
colored pigment. It segregates as a simple recessive and breeds true. 
It is of interest as being one of the few known instances in the butter- 
flies in which a single gene manifests its effect not only in the adult 
but also in the earlier stages. 

ab minor 

About 200 final instar larvae, which had been kept well supplied 
with food during their earlier stages, were allowed to feed normally for 
3 days and were then starved. About half died, the remainder wander- 
ed around their cage and eventually pupated. Normally, at 20 °C, 
the final instar larvae feeds for 5 days before pupation ( David & Gard- 
iner, 1962). 

The majority of the pupae produced butterflies, all ab minor, the 
smallest specimens having a wing span of 37 mm. male, 38 mm. 
female. 

As a control siblings to the starved larvae were kept and allowed 
to feed normally and pupate when ready. These all produced normal 
sized butterflies. 

Eggs obtained by the pairing of several of these minor were reared 
and an adequate supply of food was given throughout their life. Normal 
size specimens were produced. 

It is evident from this result that ab minor can be produced by a 
condition of the external environment, starvation, on the larvae. Never- 
theless the size of brassicae is, at least in some instances, genetically 
controlled. This will be dealt with more fully in the discussion. 

ab nigroviridescens 

About a hundred prepupae of the Cambridge stock were put in a 
desicator containing water to give 100% R.H. and kept at 20 °C. One 
reasonable specimen of nigroviridescens Rocci emerged and two par- 
tially crippled adults intermediate between normal and nigroviridescens. 


132 


GARDINER 


]. Res. Lepiil. 


No breeding was done from these nor have any similar specimens ever 
been noted in the Cambridge stock. It is considered probable, how- 
ever, that the nigroviridescens produced was a result of the saturated 
humidity of the environment. 

ab coerulea/ albinensis 

The cross between coerulea and albinensis was performed both 
ways. One brood, coerulea $ X albinensis $ being reared by the 
author and the brood from the reciprocal cross being reared by Mr. 
C. F. Rivers of the Agricultural Research Council’s Virus Research Unit. 

In both crosses the butterflies used were from a pure bred line of 
the respective recessive. 

In both crosses the several hundred individuals of the Fi were all 
normal brassicae. 

The F^ of both crosses gave a ratio very close to 9 : 3 : 3 : 1 as is 
to be expected from the independent assortment of two pairs of allelo- 
morphs. The back-cross was also performed to give a 1 : 1 : 1 : 1 
ratio. The numbers involved are shown in Table 1. 

These results show that linkage, which has not yet been shown to 
occur in butterflies (Ford 1946) is not present. Since brassicae has a 
relatively large number, 15, of chromosomes (Lorkovic, 1941) this is 
perhaps not very surprising. It would have been more interesting to 
have crossed coeridea with jauni, since both involve a change in the 
same pigment, unlike albinensis which is caused by an absence of black. 
Unfortunately by the time coerulea appeared the jauni form had been 
lost. 


Cross 

normal 

coerulea 

albinensis albinensis 
coerulea 

coerulea $> 

X 

albinensis 2 

921 

295 

304 

97 

albinensis d 

X 

coerulea $ 

695 

233 

227 

76 

coerulea/ albinensis 

X 

double heterozygote 

97 

96 

92 

95 


Table 1. The numbers of butterflies reared in the F 2 of the double re- 
cessive cross and the Fi of the back-cross. 


2 ( 2 ): 127 - 136 , 1963 


PIERIS BRASSICAE VARIATION 


133 


As might be expected the majority of the double recessives failed 
to eclose normally but, like their respective grandparents, the percentage 
of normal eclosions rises with continual breeding. 

Homeosis 

This is extremely rare in butterflies. In hrassicae specimens are 
known in which the fore wing black markings are duplicated on one 
side of the hind wing. According to Ford (1946) there is no definite 
evidence of its nature in butterflies but it has been shown to be genetic 
in Drosophila. 

During the course of hybridising Cambridge stock hrassicae with 
race cheiranthi from the Canary Islands, three homeotic specimens 
turned up in one brood of some 300 individuals, all the progeny of one 
pairing. 

The male parent was an F 3 from a back-cross of an Fi hrassicae/ 
cheiranthi hybrid to cheiranthi, while the female was an alhinensis ex- 
tracted in the F 2 of an alhinensis crossed to the Fi hrassicae/ cheiranthi 
hybrid. The cheiranthi used for the back -cross and the Fi hrassicae/ 
cheiranthi hybrids had the same father. 

All the homeotics were male with the black markings on the fore 
wing underside partially replicated on the hind wing of one side only. 

One male was mated to his sister and sib pairing was continued 
to the F;^j some 300 individuals being reared in each generation. No 
more homeotic specimens were produced. 

This is of course a negative result, but it shows that homeosis is 
not caused by a single recessive allelomorph. The most likely explana- 
tion of its occurrence, in this instance, is an upset in the genetic com- 
patability due to a slight variation between the parent genes. 

DISCUSSION 

The continuously brooded stock of hrassicae kept at the Unit of Insect 
Physiology is now thirteen years old and provides a unique opportun- 
ity for the study of variation in a butterfly. The numbers produced 
have varied from a fev^ dozen to a thousand or more per week. 

In addition to a number of known aberrations two new ones, jauni 
and alhinensis, have been produced. A third, coerulea, was only pre- 
viously known from two examples, both taken over fifty years ago. 

As was to be expected, changes in the color of the wings, coerulea 
and jauni, are recessives. It is probable that the yellow form ab flava 
Kane is also recessive and its extreme rarity is, like coerulea, due to 
failure to eclose normally. 

The failure to eclose of coerulea and alhinensis appears to be due 
to the wings sticking to the pupal case. Indeed scales can be seen to 
be left behind in the pupal cases from which specimens have succeeded 
in getting free. It is however a disability which is overcome by con- 



Left side, from top down, ab coemlea $ allotype (upperside). $ para- 
type (underside L). d holotype (underside). $ coemlea /albinensis (upper- 
side) L. Right side. Typical $ ( underside ). ab $ holotype (un- 

derside), ab minor $ (upperside). d brassicae/cheiranthi hybrid underside 
showing homeosis. 


2 ( 2 ):} 27 -] 36 , 1963 


PIERIS BRASSICAE VARIATION 


135 


tinued breeding. This failure to eclose, however, again occurs if albin- 
ensis, from a normally emerging stock is crossed to typical hrassicae 
and extracted again in the Fi>. The effect is mechanical, more marked 
in the males than in the females and is affected by temperature. It 
might be that it is a genetic effect and it would be interesting to breed 
from the crippled specimens if this were possible. Since only perfect 
images are used for breeding, there is a very strong selection towards 
normality and only recessive genes producing crippling are rapidly elim- 
inated for butterflies which cannot fly are unable to mate or oviposit. 

Although minor can be produced by starvation the size of hrassicae 
is also genetically controlled. From time to time small backward larvae 
are found in stocks of hrassicae. Attempts to breed these have so far 
been unsuccessful. Usually the larvae die. Two or three have been 
brought to pupation, the pupae being comparable in size to minor, but 
no adults emerged. 

In 1951 the wing span of the Cambridge stock hrassicae was 53 
mm. ^ and 58 mm. $ (David & Gardiner, 1952). By I960 how- 
ever this had risen to 58.5 mm. in the male, while the females had 
decreased to 57.9 mm. (David & Gardiner, 1961). 

The continuous inbreeding of the Cambridge stock has produced 
specimens with virtually no size difference between the sexes. This is 
smaller, especially in the female than the respective male and female 
sizes of 63 mm. and 76 mm. given by Frohawk (1934). It would 
appear that Frohawk had some unusually large female specimens before 
him. The author now believes that this figure may be a misprint in 
Frohawk’s book. A 76 mm. hrassicae would be a giant specimen. Wild 
specimens caught in 1943 from the author’s collection being 58 mm. 
and 63 mm, while Fi bred examples from a freshly caught wild female 
in 1957 are 58 mm. in both sexes. 

The Cambridge stock is therefore not significantly different in 
size to some wild populations, but both it and wild populations vary 
in size over a period of years. 

In P. hrassicae race cheiranthi the males are 60-65 mm. and the 
females 65-70 mm. (12 examples measured), but the wings are also 
broader so that the butterfly looks even larger. If these are crossed to 
Cambridge stock hrassicae the Fi insects are interm.ediate in size and 
in the F 2 a range of sizes are produced. Size in this instance is clearly 
under genetic control. 

The Cambridge stock hrassicae are on the whole lighter and with 
less black than wild populations. The varieties that have turned up, 
reducta Fritsch^ colliurensis Gelin and intermediates, are all those with 
a reduction of black. ^ 

The author has in his collection a female series from heavily marked 
with a trace of fasciata Kiefer to extreme colliurensis, in which the 
remaining spot is very feint. 


136 


GARDINER 


]. Res. Lepid. 


The evidence from this and also from hybrid crosses to cbeiranthi 
(which will be dealt with fully in a subsequent paper) clearly shows 
that a multifactorial effect is involved in the degree of black on the 
wings. 

SUMMARY 

‘ Two new varieties of Pieris brassicae L., coemlea and jauni are 
named and shown to be recessive in character. 

The failure to eclose in early broods of coemlea and albinensis is 
shown to be partly influenced by temperature. 

The double recessive coemlea/ albinensis has been reared and the 
two allelomorphs concerned shown not to be linked. 

Ab. nigroviridescens is considered to be probably produced by 
saturated humidity. 

The small form, minor, can be produced by starvation. Neverthe- 
less size in brassicae is also genetically controlled. 

The size of brassicae, both bred and wild, shows variation over a 
period of years. It is also believed that Frohawk’s figure of 76 mm. 
for the female wing span is a misprint. 

Homeosis is shown not to be due to a simple recessive gene. 

The Cambridge stock of brassicae shows a tendency for reduction in 
the amount of black on the wings. The evidence is that this is multi- 
factorial. 


REFERENCES 

DAVID, W.A.L. 1957. Breeding Pieris brassicae L. and Apanteles glomeratus 
L. as experimental insects. U. Pflkrankh. 64: ^12-All. 

DAVID, W. A. L., GARDINER, B.O.C. 1952. Laboratory breeding of Pieris 
brassicae L. and Apantless glomeratus L. Proc. R. ent. Soc. Lond (A) 
27: 54-56. 

— 1961. The mating behaviour of Pieris brassicae (L.) in a laboratory 

culture. Bull. ent. Res. 52: 256-280. 

— — 1962. Observations on the larvae and pupae of Pieris brassicae (L.) 

in a laboratory culture. Bull. ent. Res. 53: 417-436. 

FORD, E. B. 1946. Butterflies. Collins, London. 

FROHAWK, F. W. 1914. A Natural History of the British butterfies. 

1934. The complete book of British butterflies. Ward, Locke, 

London. 

GARDINER, B.O.C. 1962. An albino form of Pieris brassicae (Lep., 
Pieridae). Ent. Gaz. 13: 97-100. 

GRAHAM-SMITH, G. S., GRAHAM-SMITH, W. 1930. Pieris brassicae, L., 
with special reference to aberrations from Aberdeenshire. Ent. Rec. & 
J. Var. 42: 1-7. 

LORKOVIC, Z. 1941. Die Chromosomenzahlen in der Spermatogenese de 
Tagf alter. Chromosoma 2: 151T191- 


Journal of Research on the Lepidoptera 


2(2) :137-141, 1963 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
© Copyright 1963 


TYPE LOCALITIES OF THE MEGATHYMIDAE 

H. A. FREEMAN 

1605 Lewis, Garland Texas 

It is not possible for a person to make a careful systematic 
or ecological study of any group of lepidoptera unless the type local- 
ities of all of the known species and subspecies are well known. In 
recent years this information has been carefully presented in the orig- 
inal descriptions, however some of our earlier authors were not as 
careful and we have to do considerable research in order to associate 
a present habitat with their findings. 

In working with the Megathymidae I have found that there are 
several cases where no specific type locality was given or else the 
locality was entirely too general. With that in mind this paper pre- 
sents a list of these localities (Table 1) with the hope that it will make 
it easier for future workers to have the type localities at hand while 
they are doing research on this particular group of lepidoptera. 


REFERENCES 

BARNES, WILLIAM, & JAMES H. McDUNNOUGH, 1912. A review of 
the Megathymidae. Contrih. Nat. Hist. Lepid. No. Amer. 1: 1-28, pps. 
BASSETT, ANNA S., 1938. Some Georgia records of John Abbot, Naturalist. 
Auk 55: 244-254. 

BELL, ERNEST L,, 1938a. The Hesperioidea. Bull. Cheyenne Mountain Mus., 
1: 1-35. 

• — 1938b. A new genus and five new species of Neotropical Hes- 

periidae. Amer. Mus. Novitates, no. 1013: 1-11, figs. 1-6. 

BUTLER, ANDREW G., & HERBERT DRUCE, 1872. Descriptions of new 
genera and species of lepidoptera from Costa Rica. Cistula Enet. 1 : 95-118. 
DOW, ROBERT P., 1914. John Abbot of Georgia. ]ourn. New York Ent. 
Soc. 22: 65-72. 

Thanks are expressed to the National Science Foundation for research 
grants making my work on the Megathymidae possible. 


137 


138 


FREEMAN 


J. Res. LcpiJ. 


EDWARDS, W. H., 1882. Description of species of butterflies taken in 
Arizona by Jacob Doll, 1881. Papilio 2: 19-29. 

FREEMAN, H. A., 1943. Notes on and redescriptions of Megathymus yuccae 
(Bdv. & LeC.) and its sub species. Ent News 54: 211-217. 

1950. Notes on Megathymus, with description of a new species. 

Field & Lab. 18: 144-146. 

— 1952a. Notes on Megathymus yuccae (Bdv. & LeC.), with descrip- 
tion of a new subspecies. Field & Lab. 20: 29-33. 

— — ■ — ^ — - 1952b. Two new species of Megathymus. Amer. Mus. Novitates 
no. 1953: 1-9, figs. 1-13. 

— 1955. Four new species of Megathymus. Amer. Mus. Novitates, 

no. 1711: 1-20, figs. 1-34. 

1958. A revision of the genera of the Megathymidae, with the 

description of three new genera. Lepid. News. 12: 81-92. 

I960. Notes on Agathymus in Texas, and the description of a new 

species from Mexico (Megathymidae). Journ. Lepid. Soc. 14: 58-62. 

1962. A new species of Agathymus from Texas. Amer. Mus. Nov- 
itates, no. 2097, 1-7, figs. 1-6. 

HARBISON, CHARLES F., 1957. A new species of Megathymus from Baja 
California, Mexico. Trans. San Diego Soc. Nat. Hist. 12: 231-262, pis. 
18-21. 

HOLLAND, W. J., 1930. New species and varieties of North American but- 
terflies. Ann. Carnegie Mus. 19: 155-160. 

MURRAY- A ARON, E., 1942. Herbert Morrison in Mexico. Fnt. News 
53: 142-143. 

POLING, OTHO C., 1902. A new Megathymus from Arizona. Ent. News 
13: 97-98, 1 pi. 

RILEY, CHAS. V., 1876. Notes on the Yucca Borer, Megathymus yuccae 
(Bdv. & LeC.). Trans. Acad. Sci. St. Louis 3: 323-343, figs. 25-31. 

— • 1877. Additional notes on Megathymus yuccae. Trans. Acad. Sci. 

St. Louis 3: 566-568. 

SKINNER, HENRY, 1895. A new AEgiale {Megathymus). Can. Ent. 27: 179. 

1911. The larger Boreal American Hesperiidae including Eudamus, 

Erycides, Pyrrhopyge and Megathymus. Trans. Amer. Ent. Soc. 37: 169- 
209-, pi. 10. 

STALLINGS, DON B., & J. R. TURNER, 1954. Notes on Megathymus 
neumoegeni, with description of a new species. Lepid. News 8: 77-87. 

— 1956a. Description of a new subspecies of the Megathymus yuccae 

(Bdv. & LeC.) complex. Bull. So. Calif. Acad. Sci. 55: 150-152, pi. 3. 

— — ■ 1956b. Notes on Megathymus ursus, with description of a related 

new species. Lepid. News 10: 1-8. 

— — — 1957. Four new species of Megathymus. Ent. News 68: 1-17. 

— — - 1958. A review of the Megathymidae of Mexico, with a synopsis 

of the classification of the family. Lepid News 11: 113-137, 8 pis. 

— — — — i960. A new species of Agathymus and a new subspecies of Meg- 
athymus. Ent. News 71: 109-115. 

1961. A new subspecies of Agathymus mariae from Mexico. Journ. 

Lepid Soc. 15: 19-22. 

STRECKER, HERMAN, 1871. Description of a new species of AEgiale and 
notes on some other species of North American Lepidoptera. Proc. Acad. 
Nat. Sci. Philadelphia 1876: 148-153. 

Skipper from Arizona. Bull. So. Calif. Acad. Sci. 53: 75-87, pi. 2. 

TINKHAM, ERNEST R., 1954. The biology and description of a new Giant 
Skipper from Arizona. Bull. So. Calif. Acad. Sci. 53: GE-RG, pi. B. 


2(2):137-14h 


MEGATHYMIDAE 


139 


TABLE 1 

AEGIALE Felder 

1, hesperiaris (Walker), vie. Mexico City, D, F. , Mexico 
A GA THYMUS Freeman 

I. neumoegeni (Edwards), approx. 9 mi. S, Prescott, Ariz. 

Z. c a r 1 s b a d e n s i s (Stallings and Turner), Guadeloupe Mtns. , 

Carlsbad Cavern National Park, N. M, , on the Mesa at the 
head of Yucca Canyon, elevation 5470 fe et, pH 7, 9. 

3. florenceae (Stallings and Turner). Davis Mtns., Scenic 

Drive, Texas, elev.-6Z00 ft. , pH 5. 9. 

4. judithae ( Stallings and Turner), Hueco Mtns, , approx. 8 mi. 

E. Hueco, Texas, el. 5300 ft. , pH 7. 3. 

5. diabloensis Freeman, Diablo Mtns. , approx, 5 mi, W. Victoria 

Canyon, Texas, el, 5700 ft, , pH 7, 4, 

6. mcalpinei (Freeman), 5, 1 mi, N. Marathon, Texas, flats 

near foothills of Glass Mtns. , elev. 4300 ft, , pH 7. 4, 

7. chisoensis (Freeman), Chisos Basin, Texas, elev. 5400ft, pH 5. Z. 

8. juliae (Stallings and Turner), north of Zarca, Durango, Mexico, 

on highway 45 at Kim. 1317, elev. 6300 ft, 

9. hoffmanni (Freeman), Valle de Mexico, Mexico, 

10. evansi (Freeman), Ramsey Canyon, Arizona. 

II. belli (Freeman), La Bequilla, Durango, Mexico. 

IZ. aryxna (Dyar), The accepted type of this species is figured 
in the Biologia Cent. Amer, Lep, Het, III, pi, 69, fig, 4, It 
represents very well the maculation of the specimens found 
in the Santa Rita Mtns, , Santa Catalina Mtns. , on down to- 
wards Nogales, Ariz, Murray-Aaron (Ent, News 53: 143) 
states that he and Morrison collected around Nogales and on 
down to Hermosillo, Sonora, Mexico, The specimen is label- 
led N. Sonora, Mexico, Morrison, indicating that the specimen 
was collected in the general vicinity south of Nogales, Sonora, 
since Hermosillo is in the central part of the state. We do not 
know whether the specimen in question was collected on that 
trip or another that Morrison made by himself; the dates of 
their collecting trip has been lost. There is some question 
as to the status of drucei, which was named from fig, 3 in 
the Biologia by Skinner, One specimen from the Chiricahxia 
Mtns, and one from Ramsey Canyon, Huachuca Mtns, , Ariz. 
are identical with the figure of drucei. Specimens from those 
two areas deviate slightly from those from the Santa Rita 
and Santa Catalina Mtns, I would like to indicate the type 
locality of aryxna as being the western slopes of the Patagonia 
Mtns, , S. E» of Nogales, Sonora, Mexico. 

13, baueri (Stallings and Turner), Verde Hot Springs, Yavapai 

county, Arizona, elev, 4000 ft, 

14, freeman! Stallings and Turner, nr, Bagdad, Yavapai county, 

Arizona, elev. 5000 ft. 


140 


FREEMAN 


). Res. Lepid. 


15, field! Freeman, Guadalajara, Mexico, Jalisco Highway, 1 5, 

klm. 724, elev. 4400 ft, 

16, mariae (Barnes and Benjamin), Franklin Mtns, , El Paso, 

Texas, approx, elev, 3900 ft, , pH 8, 4, 

17, micheneri Stallings, Turner and Stallings, approx, 15 mi. 

S, Allende, Coahuila, Mexico, elev, 1300 ft, , pH 7, 0. 

18, Stephens! (Skinner), Mason Valley (La Puerta), San Diego 

coxinty, Calif, 

19, c o m s t o c k i (Harbison), 2 mi, N,E, San Simon, Baja Calif, , Mex, 

20, remington! (Stallings and Turner), mountains S, of Jacala, 

Hidalgo, Mexico, on highway 85, klm, 250, elev, 6000 ft, 

21, estelleae (Stallings and Turner), plains 56 mi, S,E, Reynosa, 

Mexico, near General Bravo, Nuevo Leon, elev, 400 ft, pH 7, 3. 

22, poling i (Skinner), Baboquivari Mtns,, Pima county, Arizona 

23, alliae (Stallings and Turner), 15 mi, W, Cameron, Arizona, 

along canyon of Little Colorado R, , elev, 5000 ft, 

24, indeci sa (Butler and Druce), Costa Rica 

25, rethon (Dyar), Sierra de Guerrero, Mexico 

TURNERINA Freeman 

1, mejicanus (Bell), Guanacevi, Durango, Mexico 

2, hazelae (Stallings and Turner), near Chilpancingo, Guerrero, 

Mexico, highway 95, klm. 235, elev, 2300 ft, 

MEGATHYMUS Scudder 

1, yucca e (Boisduval and LeConte) 

a, yucca e (Boisduval and LeConte), Aiken county. South 

Carolina is here designated type locality as it is the 
nearest place to Abbot's home in Scriven county, Georgia 
where he is known to have collected, and where Yucca 
grows. Yucca smalliana and Y, flaccida, 

b, buchholzi Freeman, Jupiter, Florida. Yucca glorios 

c, stallings! Freeman, Caldwell, Kansas, pH 6, 1, Yucca 

arkansana, 

d, wilsonorum Stallings and Turner, Victoria, Tamalipas, 

Mexico. Yucca treculeana and Y.carnerosana 

e, coloradensis Riley, vicinity of Colorado Springs, Colo, 

Yucca glauca, 

f, navajo Skinner, Ft, Wingate, Zuni Mtns,, New Mexico, 

Yucca baccata, 

g, arizonae Tinkham, Mountain View, Pima county, 

Arizona, Yucca elata and Y , Thornberi, 

h, martini Stallings and Turner, Little Rock, Los Ang- 

eles county, Calif, Yucca brevifolia, 

i, browni Stallings and Turner, Salina, Utah, Yucca 

harrimaniae, 

2. cofaqui (Strecker), The allotype was collected at Boca 

Grande, Florida which is here designated as the type locality. 
The exact locality of origin of the holotype is believed to 


2 ( 2 ); 1 } 7 - 141 , 1963 


MEGATHYMIDAE 


141 


be somewhere in northern Florida near Georgia, collected 
by Morrison, Yucca aloifolia, 

3, harrisi Freeman, Stone Mountain, Georgia, Y u c c a 
filamentosa. 

4, streckeri (Skinner), Type locality here designated at 

Petrified Forest area, Arizona as photos of the type 
agrees best with material from that area. Yucca 
Standleyi,. probably, 

5, texanus, Barnes and McDunnough 

a, texanus Barnes and McDxinnough, 2d'cf,29$ collected 

by Jacob Boll at Dallas and San Antonio, Texas, To 
this collector Dallas was North Texas and San Antonio 
was South Texas, Kerrville, Texas is here desig- 
nated at the type locality by reason of being the 
nearest known’place to the above where texanus 
may be found. Yucca glauca, associate, 

b, leussleri Holland, Sand Hills near Valentine, Neb- 

raska, Yucca glauca, probable, 

6, ursus Poling, Santa Catalina Mtns, , west of Redington, 

Pima county, Arizona, pH6.1. Yucca Schottii. 

7, viola e Stallings and Turner, Carlsbad Cavern National 

Park, New Mexico, elev, 4700 ft, , pH 7, 5, Yucca Torreyi. 

8, beulahae Stallings and Turner, near Ixmiquilpan, Hidalgo, 

Mexico, highway 85, klm, 176, elev, 5700 ft. Agave ? near 
Schottii, 

STALLINGSLA Freeman 

1, smithi (Druce), Amula, Guerrero, Mexico, approx, 3000 ft, 

Manfreda maculata, probable, 

2, rhac.ulosus (Freeman), 2 mi, S, Kingville, Texas, elev, 

100 ft,, pH 7. 0, Manfred maculosa. 


142 


FREEMAN 


/. Res. Lepid. 


BIOGRAPHICAL SKETCHES 

FREEMAN, HUGH AVERY 

[1605 Lewis Drive, Garland, Texas] 

Born; Conway, Arkansas, Oct* 7, 1912 

Married; 1939, 3 children 

A„ B, ; Hendrix College, 1936 

M. S, ; Southern Methodist University, 1938 

Teacher; High school Texas, 1938-48 

Instructor; Biology, Southern Methodist University, 1948-51 
Teacher; High school, Dallas, 1951 
Interests; Started collecting lepidoptera in 1928 
Began specializing in the hesperi - 
oidea in 1938, In 194Z started in- 
tensive study of the Megathymidae, 
which is still going on. Also inter- 
ested in the study of metal marks 
and hair streaks. Has published 44 
articles on lepidoptera, most of 
which were upon the hesperioidea. 



Journal of Research on the Lepidoptera 


2 ( 2 ) : 143 - 150 , 1963 


1140 W. Orange Grove Ave., Arcaiia, CaUforms, US. A. 
© Copyrigbi 1^63 


THE DISTRIBUTION OF 
AN ENDEMIC BUTTERFLY 
LYCAENA HERMES 

FRED THORNE 

1360 Merritt Drive 
El Cajon, California 


One of the most interesting of the endemic butterflies is 
Lycaena hermes (Edw.). Its known range extends fifty miles north 
of the Mexican border almost to Failbrook, San Diego County, Cali- 
fornia, and south of the border aimos; one hundred miles to a point 
eighteen miles south of Santo Tomas, Baja, California, Mexico. In 
San Diego County it ranges inland to Pine alley, about forty miles 
from the Pacific Ocean. 

Older literature and check-lists tended to overstate the range, i. e., 
California and Nevada, whereas all recently published information 
understates the range. In the light of present knowledge, the area 
occupied may be as large as the state of Connecticut. However, within 
this range its distribution is limited to pockets where the larval 
food plant occurs, so that the total area where the insect acually flies 
is probably not more than a fraction of one percent of the maximum 
area. Such limited distribution within a given range is not unique 
with endemics, being a common occurence among plants and animals. 

Colonies of the Hermes Copper are closely confined to the vicinity 
of the host plant, Rhamnus crocea Nutt. Extensive collecting for over 
thirty years has failed to produce specimens beyond a short distance 
from the larval food plant. There is no observable tendency to migrate, 
to "hilltop,” or otherwise to stray from these colonies, although there 
must be some inter-colony movement, probably by the males. Popula- 
tions within the known range therefore depend on the distribution of 
the host plant, and there is. certainly nothing novel about this among 
insects. 

It is very difficult to analyse the complex factors- which determine 
why a certain plant has been successful in a 'given spot, and why 
it has been able to out-compete all other plants for this particular 
place in the sun. In the case of Rhamnus crocea, the only consistent 
requirement seems to, be a well drained soil of better than average 
depth, yet not deep enough to support trees. Such soils occur along 
canyon bottoms and on hillsides with a northern exposure; therefore, 
it is in these, situations that hemes is -generally found. 


143 


144 


THORNE 


). Res. LepiJ. 


A notable point about hermes is that the host plant extends well 
beyond the range of this insect, Rhamnus crocea, in one form or another, 
extends to Mt. Diablo in the coastal ranges of California; along the 
foothills of the Sierra Nevada; on the Channel (Santa Barbara) Islands; 
and even into the Mojave Desert. Yet within its range in San Diego 
County, hermes tolerates greater climatic extremes • than it would en- 
counter in some of the contiguous coastal areas open to colonization, 
and in insular areas which might have been available at one time. 

Freezing winters with snow are normal in Pine Valley at an 
elevation of 3800 feet, while summer temperatures of 105 °F. are not 
uncommon in some of the foothill localities. To borrow an appropriate 
sentence from Flovanitz (1963), said of the distribution of Argynnis 
idalia, "The biological reasons for this restricted distributional range 
are not known." Biologists will appreciate how often this same con- 
clusion must be drawn. 

Nelson (1921) recognized a San Diegan Faunal District in north- 
western Baja California which roughly corresponds with the area 
occupied by hermes in that Mexican state, but did not define the 
northern limits of this life zone in Southern California. A few species 
of plants appear to be restricted to about the area inhabited by 
hermes, but there is little to suggest, at least north of the border, that 
the area should be segregated from the Upper and Lower Sonoran 
life zones usually assigned to it. Nor has northwestern Baja California 
been shown to be an originator of new species, since the coastal fauna 
shows a close affinity to that of coastal Southern California, and the 
mountain fauna is simply an extension of the southern Sierran with 
the exception of a few instrusions (Rindge, 1948; Powell, 1958; Pat- 
terson and Powell, 1959; Truxal, I960). 

L. hermes is in a good state of balance in its environment. The 
season of emergence for the adults is very dependable, as is their 
presence every year in their select habitats. There is no wide fluctuation 
in numbers from year to year, although the current prolonged drought 
has reduced the populations in common with nearly all Lepidoptera. 
It would be difficult to believe that it is not autochthonous. Fossil 
evidence of insect distribution is so limited that it will probably 
never be known whether hermes ranged over a wider territory than 
now. It is an insect which seems to exhibit stability due to long occupa- 
tion of its present habitat, yet it is difficult in the light of other biologi- 
cal evidence to view the present range as a refugium of some sort. 

Concerning the genus Lycaena, Clench (1961) has stated, "A 
curious and possibly quite ancient genus, strongly developed in both 
the Palaearctic and the Nearctic regions, with a small handful of 
outliers — one in Guatemala, one in South Africa and several, most 
perplexing, in New Zealand.’’ Perhaps this offers a clue, yet careful 
studies by Klots (1936) and Freeman (1936) have indicated that 


2 ( 2 ): 143 - 1 ^ 0 , 1963 


DISTRIBUTION LYCAENA HERMES 


145 


hermes has affinities with other North American members of the 
genus. Kiots says, "Evidently hermes is, structurally at least, far closer 
to gorgon and heteronea than to its tailed Nearctic congeners." Free- 
man placed hermes in his xanthoides group on the basis of the genitalia, 
together with dione, rubidus, and editha. 

Another interesting thing about hermes is the use of a species of 
Rhamnaceae as the larval host. Most congeners feed on Polygonaceae 
(Eriogonum, Rumex, Polygomim)\ some of Rosaceae {Potentilla) ] 
some on Saxifragaceae {Ribes)\ and some on Ericaceae {Vaccinium), 
but this use of a Rhamnaceous plant is believed to be unique for 
the genus (Davenport and Dethier, 1937). The commonest source 
of nectar for the imagines is Eriogonum fasciculatum Benth., one of 
the Polygonaceae, and this plant is almost invariably present. Could 
this in some ancient time have served as the larval host? 

One might reason that the failure of hermes to invade large areas 
which appear to be open to colonization might be due to prior occupa- 
tion of particular ecological niches by another species (competitive 
exclusion). The unusual food plant which is not used in other areas 
by any member of the genus would seem to rule this out. Or, if one 
wishes to consider hermes as the victim of some complex, predator- 
parasite relationship which grew up around one of the congeners 
and favored it over hermes, Lycaena {T harsalea) arota would be the 
most logical candidate. This species appears to be at the extreme 
southern limit of its range in San Diego County as evidenced by the 
few specimens ever taken. Since it feeds on Ribes, there is no direct 
competition, and the hypothesis becomes even more dubious from 
the fact that hermes lives sympatrically, or at least there are zones of 
contact along watercourses with Lycaena helloides and L. xanthoides. 
These should reasonably be expected to furnish whatever environ- 
mental pressures that arota might. 

There is rather general belief that hermes is in a last ditch struggle 
for survival in San Diego County. This isn’t true! Colonies have 
survived in areas that have been overrun with houses for many 
years; in areas being grazed by livestock; in areas being farmed 
(avocado orchards); and in areas which have been burned-over with 
some frequency. The map, Fig. 1, shows the wide distribution of 
known colonies which should ensure survival for the foreseeable 
future. This map should not be regarded as a complete record of 
distribution, since accessibility by road has been the main factor in 
locating colonies. 

The insect has been beautifully illustrated in Comstock’s "Butter- 
flies of California” and his poetic description of the butterfly in 
nature is worth repeating. He says, "It is a fascinating little sprite as 
it darts about in the sunlight, or sports is showy colors while balanced 
on a tuft of wild buckwheat." As for the flight period, my earliest 


146 


THORNE 


J. Res. LepiJ. 


record is May 20, 1934, and the latest is July 20 at Alpine by George 
Field. Records from several hundred captures show peak flight about 
June 20, but the best time to collect males is about June 10, and 
for the females about June 20. Field captures show a large preponder- 
ance of males (85%), but this a probably a false indication of the 
actual sex ratio because of the mo’-e retiring habits of the females, and 
because of their tendency to flee directly from the place of disturbance 
so that they are quickly lost to sight. However, the percentage of 
females increases late in the flight period in common with many 
butterflies. The males practice territorialism, but are not very aggre- 
sive about it. They will patrol a section of flyway, or watch it from 
a vantage point, often on the host plant, but from any suitable perch. 
Both sexes visit flowers avidly, and the blossoms of Eriogonum fasci- 
culatum supply the bulk of nectar. 

The species is single-brooded and spends about two-thirds of its 
life in the egg stage. It aestivates and hibernates in the ovum, and the 
hatching of one egg was observed under field conditions on March 16. 
Mature larvae were recovered by beating the host plant on May 24 
near Lyons Peak, where the season is delayed due to elevation. The 
egg, mature larva, and pupa have been illustrated by Comstock and 
Dammers ( 1935). Females oviposit readily in captivity, but unless 
the ova are kept on a living plant, they fail to hatch. The reason for 
this is not known, but may simply be dessication. Nothing in this 
life history sets hermes apart in any remarkable way from other mem- 
bers of the genus, although several are multivoltine. 

As a generalization, most endemics are univoltine (consider the 
alpine relicts) but there are numerous exceptions. A good example is 
Strymon avalona Wright, another interesting endemic, which is con- 
fined to Santa Catalina Island off the southern California coast despite 
the fact that its food plant, Hosackia (Lotus) argophylla Gray, is more 
widespread. This insect has a succession of broods. The biological 
reasons for its restricted distributional range are not known, but are 
easier to deal with than is the case with hermes, since insular, alpine, 
acid bog, or other types of endemism offer the biologist some 
solace during the brief instant in evolutionary time that he is around 
to observe distribution. 

Populations of the Hermes Copper in each colony are not great, 
probably numbering in the hundreds. Six sample counts taken at 
random from field notes for 1955 to 1959 show the capture of 69 
specimens in 405 minutes, about 6 minutes per catch. Any day in 
which 50 specimens are taken can be regarded as exceptional. It is 
entirely fair to regard the insect as "not uncommon" as expressed by 
Clench (1961) — in fact, it falls comfortably into Clark’s (1932) 
standard of "abundant” (where fifteen or more can be taken in an 
average day ) . 


2(2):14}-n0, 196} 


DISTRIBUTION LYCAENA HERMES 


147 


, PAL OMA R M TN. 
ST ATE PARK 


O 



Fig. 1. Map showing the distribution of Lycaena hermes in San Diego 
county, California. . i 


148 


THORNE 


). Res. Lepid. 


It has been stated previously that there must be some inter-colony 
movement. The basis for this is that differences among the populations 
are not readily observable, if indeed any exist. It must be admitted 
that no effort has been made to comoare adequate series from 
different colonies to see if any segregation is evident. This vv^ould be an 
interesting study, but until it is made, it seems best to assume that 
gene flow throughout the entire range is adequate to prevent segregation. 

If this is true, then the gene pool has some magnitude. Nevertheless, 
it seems likely that hermes is a conservative species, homozygous for 
many of its characters, variability being restricted by a close adaptation 
to a narrow environment. Barriers to spread appear to be intrinsic, 
that is, because of the inherited behavior patterns, the imagines "choose 
to remain” within very limited areas despite their ability to fly else- 
where, and presumably, to occupy larger territories (see Ehrlich, 1961). 
The chromosome number has not been published yet (N=24 for most 
species of the genus (Maeki and Remington, I960) but this infor- 
mation is expected to be available soon. 

Perhaps the distinctive facies and unusual food plant of hermes 
represent a genetic breakthrough which will result in time in a more 
widespread and successful species. Only time will tell whether the 
insect has retreated into a final refugium, the nature of which is not 
too evident, or whether it has the genetic resources eventually to 
expand its habitats. 

That such expansion is possible has been demonstrated in a 
spectacular way by Paratrytone melane melane (Edw.) which was 
not recorded from San Diego County prior to 1941. Wright (1930) 
said, "Further collecting in wooded areas of the county may produce 
this species.” Gunder (1930), in his now classic checklist of the 
butterflies of Los Angeles County wrote, "Never seemingly abundant 
in one locality, but may be had, several at a time each year.” Rindge 
(1948) recorded this insect from southern Lower California, but 
this is now regarded as a distinct and undescribed sub-species (Mac 
Neill, 1962). 

I first encountered this skipper on July 20, 1941, when two speci- 
mens were collected near El Cajon, California. Visits to the same 
spot July 23, 24, 25 and August 2 and 3 resulted in fifteen more 
specimens, and show my interest in what I thought was a once-in-a- 
lifetime chance. Other collectors also reported finding this species for 
the first time in San Diego that year. On October 5, 1941, a specimen 
was taken in the desert at Mason Valley, San Diego County. Since 
then, quite contrary to my expectations, this insect has become one 
of the very common skippers in my garden, flying from February 
to December. Powell (1958) records it from northwestern Baja, 
California in what I believe to be a further extension of this same 
population explosion. 


2(2):l4}-n0, 196) 


DISTRIBUTION LYCAENA HERMES 


149 


This is an example of an insect which suddenly expanded into 
and occupied contiguous areas which are evidently well suited to 
it. It is not the purpose of this article to discuss this interesting 
phenomenon, but simply to point out that extensions of range are 
possible, and that it may be the privilege of the lepidopterist to see 
it happen in his own yard. 


SUMMARY 

The known range of Lycaena hermes (Edw. ) extends from fifty 
miles north of the Mexican border in San Diego County to one 
hundred miles south of the border in Baja California. The insect occurs 
in colonies around the foodplant Rhamnus crocea, but has failed to 
invade other areas that appear suitable for reasons that are not known. 
The life history and field behavior are not unusual. The species is 
believed to be autochthonous and conservative, perhaps in a final 
refugium, but spread to other areas is possible, as has been demon- 
strated by Paratrytone melane (Edw.) 

REFERENCES 

BROWN, F. MARTIN 1962. Notes about the types of some species of 
butterflies described by William Henry Edwards. Ent. News. 73 (10): 
265-268 (pg. 267). 

CLARK, AUSTIN H. 1932. Butterflies of the District of Columbia. Smith- 
sonian Inst. U. S. Nat. Mus. Bull. 137, Wash. D. C. (pg. 25), 
CLENCH, HARRY K. 1961. In, How to know the butterflies. Ehrlich and 
Ehrlich. Brown Co., Dubuque, Iowa (pg. 22). 

COMSTOCK, JOHN A. 1927. Butterflies of California. Los Angeles, Calif. 
(pg. 172 and PI 51, Figs. 9-11). 

COMSTOCK, JOHN A. and CHARLES M. DAMMERS 1935. Notes on the 
early stages of three butterflies and six moths from California. Bull. So. 
Calif. Acad. Sci. 34 (2); 120-141. (pp. 124-126). 

DAVENPORT, D and V. G. DETHIER 1937. Bibliography of the described 
life histories of Rhopalocera of America north of Mexico 1889-1937. 
Ent. Amer. 17 (New series) (4); 155-196. (pp. 174-175). 

DRAUDT, Dr. M. 1924. In Seitz, Macrolepidoptera of the world Vol. 5. 

American Rhopalocera. Stuttgart, Germany, (p. 812). 

EDWARDS, WILLIAM HENRY 1870. Description of new species of Lepi- 
doptera found within the United States. Trans. Amer. Ent. Soc. 3:21 
original description of Chrysophanus hermes.) 

EHRLICH, PAUL R. 1961. Instrinsic barriers to dispersal in checkerspot 
butterfly. Science 134, No. 3472: 108-109. 

FREEMAN, T. N. 1936. Notes on the specific grouping of the genus Lycaena 
( Lepidoptera ) . Can. Ent. 68: 277-279. 

GUNDER, J. D. 1930. Butterflies of Los Angeles County, California. Bull. 

So. Calif. Acad. Sci. 29 (2): 1-59 (on page 58). 

HOLLAND, W. J. 1931. Butterfly Book (Revised ed.), Doubleday and Co.. 
New York, (on page 247). 

HOVANITZ, WILLIAM 1963. Geographical distribution and variation of 
the genus Argynnis. J. Res. Lepid. 1 (2); 117-123 (on page 123). 
JEPSON, WILLIS LINN 1925. Manual of the flowering plants of California. 
Univ. of Calif. Press, Berkeley, Calif, (on pages 614-615). 


150 


THORNE 


}. Res. Lepid. 


KLOTS, ALEXANDER B. 1936. The interrelationship of the species of the 
genus Lycaena Eabricius (Lepid., Lycaenidae), Bull. Brooklyn Ent. Soc. 
31 (4); 154-171. (on page 164). 

MacNEILL, C. DON 1962. A preliminary report of the Hesperiidae of Baja 
California. Proc. Calif. Acad. Sci., 4th series, 30 (5): 91-116. 

MAEKI, KODO and CHARLES L. REMINGTON I960. Studies of the 
chromosomes of North American Rhopalocera, Part 3. /• Lep. Soc. 14 (2) : 
127-147. 

MUNZ, PHILIP A. 1935. Manual of Southern California Botany, Claremont 
Colleges, Claremont, Calif, (on page 299). 

NELSON, EDWARD W. 1921. Lower California and its natural resources. 
Mem. Nat. Acad. Sci., Wash., D. C. 16: 1-194. (on page 118). 

PATTERSON, DONALD and JERRY A. POWELL 1959. Lepidoptera col- 
lecting in the Sierra San Pedro Martir, Baja California J. Lep. Soc. 13: 
229-235. 

POWELL, JERRY A. 1958. Additions to the knowledge of the butterfly fauna 
of Baja California Norte. Lep. News 12: 26-32. 

RINDGE, FREDERICK H. 1948. Contributions toward a knowledge of the 
insect fauna of Lower California. No. 8 Lepidoptera; Rhopalocera. Proc. 
Calif. Acad. Sci., 24: 289-311. 

TILDEN, J. W. 1955. A revision of Tharsalea Scud. (s. str.) with descrip- 
tion of a new subspecies (Lepid., Lyc. ) Bull. So. Calif. Acad. Sci. 54 
(2): 67-77. 

TRUXALL, FRED S. I960. Symposium: Biogeography of Baja California 
and adjacent seas. The entomofauna with special reference to its origins 
and affinities. Syst. Zoo. 9 (1-4): 165-170. 

WRIGHT, WILLIAM GREENWOOD 1906. Butterflies of the West Coast. 
Second ed. San Bernardino, Calif. 

WRIGHT, WILLIAM S. 1930. Annotated list of the butterflies of San Diego 
County, California. Trans. San Diego Soc. Nat. Hist. 6 (1): 1-40 (on 
pages 25, 26 and 40). 


Journal of- Research on the Lepidoptera 


2 ( 2 ) : 151 - 160 , 1963 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
© Copyright 


CALLOPHRYS (LYCAENIDAE) FROM 
THE PACIFIC NORTHWEST 

HARRY K. CLENCH 

Carnegie Museum, Pittsburgh, 13, Pa. 

Over two years ago Mr. E. J. Newcomer, of Yakima, Wash- 
ington, sent for examination and identification a series of Callophrys 
{Callophrys) taken near his home. He subsequently augmented this 
lot by other specimens, some of- which were also taken locally while 
others were from farther afield in the state. More recently Mr. David 
L. Bauer, of Bijou, California, has sent for study additional specimens 
of this subgenus from both Washington and Oregon. The present 
paper is concerned chiefly with this assembled material. My best 
thanks are due both these gentlemen for the opportunity to study 
and report on this now sizable and instructive collection from a region 
whence before Callophrys had been all but unknown. 

Callophrys is in several ways a very difficult subgenus. Problems 
of discriminating species occur in several regions, notably (but by no 
means exclusively) that under discussion in this paper. An even greater 
problem exists in the group as a whole: the correct association in 
polytypic species of the various named entities. Six species are cur- 
rently recognized (Clench, 1961) in the subgenus {dtimetorurn- Bois- 
duval, corns tocki Henne, apama Edwards, af finis Edwards, viridis 
Edwards and sheridanii Edwards ) . a number which is probably too 
large. On the other hand^ we know that there must be at least two 
species for two occur sympatrically in many places: sheridanii and 
apama in Colorado; sheridanii and af finis in the Yellowstone area 
and near Brewster, Washington, sheridanii and dumetornm near 
Yakima, Washington; viridis and dumetornm in the San Francisco Bay 
area. But throughout their known ranges apama and affinis do not; 
nor do affinis and dumetornm; apama and dumetorum; sheridanii and 
viridis; or affinis and viridis. 

It is tempting, and it may possibly be correct, to associate sheridanii 
and viridis as one species; dumetorum, comstocki (which is sympatric 
with no other), apama and affinis as another. This arrangement would 
satisfy nicely the conditions of sympatry, but it creates other problems; 
and besides, it is sheer speculation. 

The traditional method of resolving such difficulties is by study 
of the male genitalia. Barnes and Benjamin (1923), the first authors 
to revise the group, did not make use of these structures nor did I 


151 


152 


CLENCH 


). Res. Lepid. 


in my revision of some years ago (Clench, 1944). The present occasion 
seemed an excellent opportunity to fill this long-standing desideratum. 
Preparations were accordingly made of all of the major and most of 
the minor entities, usually of several individuals each, and these prep- 
arations subjected to study. Preliminary examination failed to disclose 
any qualitative differences but did reveal the existence of considerable 
variation in relative lengths of different structures. A series of measure- 
ments was then made and these carefully and elaborately analyzed 
individually, paired in correlations, as ratios and by other more esoteric 
means, which it is pointless to describe as the results of the whole 
inquiry were thoroughly, discouragingly negative: the male genitalia of 
Callophrys s.s. show considerable individual variability but give no 
evidence whatever of variation useful for discriminating species. We 
are still where we were before. 

Three species of Callophrys s.s. are now known from the Pacific 
Northwest^ one here recorded for the first time. They epitomize the 
difficulties noted just above and it would be fatuous to presume that 
the results here given represent the final word on the solution of those 
difficulties. Decisions on relationships had to be made and I have 
tried to make them as accurately as possible. 

Callophrys {Callophrys) af finis washingtonia Clench 

Known from Alta Lake (type locality) and Brewster, Washington 
and from Summerland, Osoyoos, Penticton and Keremeos, British 
Columbia, the latter records in the Canadian National collection. 

Size rather large (about as in dumetorum perplexa or apama)\ 
ground color above greyish with a distinct brownish tinge and in the 
female with some discal fulvous (occasionally in the male as well); 
underside apple green, a little less yellowish than in nominate affinis 
but decidely yellower than other members of the genus; on the fore- 
wing the green extends posteriorly to about CU 2 , thus covering most 
of the wing; postmedian spot row completely absent or at most repre- 
sented by faint, hardly visible traces. Base of fore wing costa below 
pale grey. 

Callophrys {Callophrys) dumetorum Boisduval 

When Mr. Newcomer sent his first series of Callophrys he wrote 
that he was interested to learn whether they represented one species 
or two, for some were immaculate below like affinis, others well 
marked, more or less as in sheridanii. Study showed that all of this 
variability actually was comprised in the one new subspecies of 
sheridanii described below. There was a surprise for us both, how- 
ever, that came to light only when the specimens were being studied 
after spreading: a single fresh male of dumetorum from Satus Creek, 
2000 ft., Yakima Co.^ Washington, taken in 16 May I960. When I 
wrote to him about this discovery and described its' differentiating 
characters, Mr. Newcomer checked through his material and found 
an additional specimen from the same locality taken I 6 May 1959. 
Returning to the same area in the spring of 1961 he was able to 


2(2):151-I60, 1963 


CALLOPHRYS FROM NORTHWEST 


153 


take several additional specimens most of which he sent on for 
study: Kussdhi Creek, off Sams Creek, 2000 ft., Yakima Co., 24 
May 1961 (1 female), and 2 June 1961 (1 male); Ski area near 
Sams Pass, 3700 ft., Klickitat Co., 2 June 1961 (1 female). In the 
material sent by Mr. Bauer were two males of this species from 
Shelton, Mason Co., Washington, 2 May 1958. All these specimens 
represent a considerable northward extension of the known range of 
dumetofum (cf. Clench, 1961; 210). 

Three regional lists relating to the area under consideration in this 
paper have had to be ignored: Bowman (1919) on Alberta; Blackmore 
(1927) on British Columbia; and Leighton (1946) on Washington state. 
Each lists dumetomm and no other Callophrys s. s. (except Leighton, who 
also lists dumetomm perplexa and sheridanii) , misdeterminations (partial 
or complete) rather typical of the trouble these butterflies have given to 
systematists. 

Males [Description refers only to Washington material.] are uniform 
grey above with a very slight brownish cast. Females above are 
largely dark fulvous, shading to brownish fuscous in the basal third 
of both wings, along costa, termen and (more broadly) apex of fore 
wing as well as very narrowly along termen of hind wing. The fringe 
is greyish white (markedly duller than the fringe of either af finis or 
dumetomm), darker basad. On the under surface of both sexes the 
inner marginal grey of fore wing extends costad usually to M 3 ; the 
postmedian line of this wing is tolerably well developed from Cuq to 
Ml or R 4 , but often rather faint, occasionally even obsolete (though 
a trace of the dark basal part of the line usually persists ) . The post- 
median line of the hind wing is highly variable in its expression, rang- 
ing from nearly complete (bars in ail interspaces from Sc to inner 
margin) to nearly absent (slight whitish bars in Sc - Rs and Cui - 
CU 2 ); base of fore wing costa below usually fulvous. Size rather small: 
about the same as s. sheridanii. 

The subspecies of dumetomm have never been worked out adequa- 
tely. Even the range of the species is imperfectly known. For the 
present these Washington specimens seem best left under nominate 
dumetomm with which they agree far more closely than with d. 
perplexa Barnes and Benjamin from the lowlands of southern Cali- 
fornia. 

Callophrys {Callophrys) sheridanii neoperplexa Barnes and Benjamin 

The upper surfaces of both sexes are uniform grey with no 
tinge of brownish or fulvous. Base of fore wing costa below pale 
grey. Under surface green, on fore wing extending posteriorly to 
Cu 2 ; postmedian lines well developed on both wings, usually more 
or less continuous (the component white bars conjoined), but rather 
thin and lacking the black basal edging characteristic of s. sheridanii. 
Size small. 

In the Pacific Northwest known only from Brewester, Washington 
where the late J. C. Hopfinger used to take it rather frequently. Until 


154 


CLENCH 


). Res. Lepid. 


the receipt of Mr. Newcomer’s material it was not realized just how 
unusual this record is. See the discussion below. 

Callophrys ( Callophrys ) sheridanii newcomeri new subspecies 

The upper surfaces of both sexes are uniform grey, unrelieved by 
any brownish tint or fulvous, much as in the other sheridanii sub- 
species. Base of fore wing costa below, pale grey. It differs chiefly in 
the strong tendency to reduction of the postmiedian line of the under- 
side, continuing the trend away from s. sheridanii that is evident in 
s. neoperplexa Barnes and Benjamin. This is especially marked on 
the fore wing where most specimens show either no trace of the 
line at all or only a few faint bars; occasionally it may be fully 
present but is then only faintly developed. In only one specimen, a 
female from Ft. Simcoe, is it fully present and strongly developed. 
The row on the hind wing is rarely completely absent (a female 
from Mill Creek), but almost always lacks several bars at least, 
especially those in Rs - Mi and M 3 - Cui. On the fore wing upper 
side of the Mill Creek series there is a definite, though faint, pale 
grey patch, strong at cell-end and fading rapidly distad. 

Holotype, male, and 5 male paratypes, all Mill Creek, 1800 ft., 
Yakima Co., Washington, 29 March 1961 (E. J. Newcomer) the 
holotype including male genitalia slide C-791; 5 male and 7 female 
paratypes, Ft. Simcoe, 1200 ft., Yakima Co., Washington, 6 April I960 
(E. J. Newcomer). 

Holotype and most of the paratypes, C. M. Ent. type series no. 
478. Some paratypes are being returned to Mr Newcomer. 

Remarks In addition to the type series, the following material 
has been seen: Mt. Spokane, 5000 ft., Spokane Co., Washington, 27 
June I960 (E. J. Newcomer), 3 males and 2 females; Blue Mts., 
4800 ft., Columbia Co., Washington, 17 June 1961 (E. J. Newcomer), 
2 males; Lonerock, Gilliam Co., Oregon, 7 June 1961 (D. L. Bauer), 
2 males and 2 females (doubtfully typical). 

The Canadian National Collection has specimens from Waterton 
Lakes Park, Alberta, and from Okanagan Landing and Vernon, 
British Columbia, all of which I believe are referable to newcomeri, 
though I saw them but briefly while on a visit in 1956 and made only 
a few notes on their peculiarities. 

The Oregon population and the four from Washington all show 
a certain amount of independent differentiation. The greatest departure 
is found in the Lonerock series where females show a definite fulvous 
tinge above, unique in the species, and the under surface (which is 
more uniform in appearance than in any of the Washington popula- 
tions) has the postmedian line on both wings practically obsolete save 
for the portion posterior to Cui on the hind wing. Should these dif- 
ferences hold true in a larger series it may be advisable to separate 
this under another subspecific name. 


2(2):IU-160, 196} 


CALLOPHRYS FROM NORTHWEST 


155 


The fresh series of males from Mill Creek shows a faint, small, 
pale discal patch on the fore wing above as noted above; this also 
occurs in some of the males from Ft. Simcoe, though more weakly. 
The Mill Creek series also averages somewhat larger than the others. 

The specimens from Mt. Spokane are somewhat smaller, tend to 
be somewhat darker below and have the postmedian line of the 
fore wing below absent completely. 

Several points concerning sheridanii in the Northwest merit 
further discussion. These points may be grouped under two broad 
headings: geographic variation and climatic adaptation. 

Geographic variation. The differences between sheridanii popula- 
tions from place to place are of a rather unusual nature. The most 
conspicuous aspect of this is the isolated occurrence of s. neoperplexa 
at Brewster, Washington, remote from the nearest other populations of 
that subspecies in southwestern Montana and with much of the 
intervening area occupied, apparently by s. newcomeri ( see fig. 1 ) . 
It is not yet known whether this Brewster neoperplexa represents an 
enclave entirely cut off from other neoperplexa populations or a 

peninsula-like intrusion, perhaps along river valleys. 

The population of neivcomeri at Lonerock, Oregon, differs more 
from any of the Washington populations than these do from each 
other, rather surprizing in view of its close geographic proximity. 
Lonerock and Ft. Simcoe, for example, are much closer than Ft. 
Simcoe is to Mt. Spokane; and both Lonerock and the Blue Mts. locality 
in Columbia Co., Washington, are on the same (northwest) slope 
of the same mountain range. In addition to its greater differentiation, 
the Lonerock series also seems less individually variable. Again it 

should be pointed out that the small size of the series makes these 
observations tentative and uncertain. 

In Washington the known populations of newcomeri show a 
definite but rather low level of interpopulation differentiation as 

already discussed; and within each population a rather high level of 
individual variability. The interpopulational differences seem to bear 
little relation to the geographic distances separating the populations, 
the Mill Creek and Ft. Simcoe populations, for example, being only 
a little less different from one another than either is from the Mt. 
Spokane series. 

There is evident here a definite hierarchy of differentiation: the 
greatest, that between newcomeri and neoperplexa; next, in neivcomeri, 
between the Washington populations collectively and the one popula- 
tion from Oregon; and finally, the slight differentiation between the 
Washington populations. In a general way this hierarchy undoubtedly 
reflects the past history of the species in this region and it is a 
temptation to draw on it as well as on some of the other data on 
variation given above in speculation on the past events that may 


156 


CLENCH 


I. Res. Lct>id. 


liave led to the situation as it is today. Until larger series can be 
obtained, however, the data basic to such speculation would be far 
too shaky. My repeated emphasis of this point is in no way intended 
as a slight to the much appreciated efforts of Mr. Newcomer and 
Mr. Bauer; their collections are by no means small as series go in 
Callophrys and, further, they were not collecting with statistical needs 
in mind. Statistics is a notoriously avaricious taskmaster. 

Climatic adaptation. The various factors which together make 
up what we call climate exert a strong control over the distribution 
of Lepidoptera as they do for many other groups of organisms. Indeed, 
we may imagine the range of a species of butterfly or moth as an 
area bounded by not one but a number of lines, each one representing 
a limiting value, for that species, of some particular climatic factor. 
Such a notion, evidently, is a great oversimplification but the principle 
is valid and useful. 

We may touch on the problem only briefly and incompletely 
here in connection with C. sheridanii. Two types of climatic responses 
may be discerned: (1) responses to particular factors singly, and 
( 2 ) responses to two or more jointly ( correlated responses ) . The 
data used are weather bureau mean figures as recorded, for example, 
in Climate and Man (U.S.D.A. Yearbook of Agriculture, 1941). On 
no account is it to be imagined that the insects are necessarily respond- 
ing directly to these variables as such. It is merely that they appear 
to function as indices more or less closely correlated with whatever 
factors may actually be responsible, factors which themselves may be 
quite inaccessible for analysis over a large area. 

Three variables were selected for study: mean January temperature, 
m.ean July temperature, and mean annual precipitation. Values of these 
were tabulated for all the known sheridanii localities for which they 
were available, as given in Table 1. By inspection we may establish 
the approximate limits for each of these; 

mean January: all records fall between 14° and 32 °F 
mean July: all records fall between 57° and 76° F 
ann. precip.: all records fall between 10 and 25 inches 
The map ( fig. 1 ) shows the results of applying these limits over the 
northern part of the range of sheridanii: all the shaded areas lie out- 
side them and hence are presumably unavailable to sheridanii. 

Obviously these three factors are not sufficient to account for the 
whole of the present range of the species. A large part of Montana 
(east of the mountains) and of South Dakota, for example, fall with- 
in these limits, yet sheridanii does not occur there. It is quite probable 
that variables other than these three are, at least in part, responsible. 
Yet without extending observation beyond them the distributional 
limits of sheridanii could be approximated still more closely by the 
use of correlated responses. 


2(2):I)I-I60, /y6.i 


CALLOPHRYS FROM NORTHWEST 


157 


TABLE 1 

Mean temperature (® P) for July and January tod mean annual preclpata- 
tlon (inches) for localities where Callophrys (Callophrys) sheridanil 
has been taken. 


Station 

Locality 

mean 

Jan. 

mean 

Jul. 

meui ani 

precip, 

1 

Wash.: 

Ft. Simcoe 

28.4 

75.2 

12.29 

2 


Blue Mts, (Columbia Co.) 

28(ca.) 

68(ca.) 

? 

3 


Mt. Spokane, 5000 ft. 

17.2 

58.7 

22(ca.)^ 

4 


Brewster 

27.6 

73.8 

10.66 

5 

Oregon; 

Lonerock 

31.7 

62.0 

15.442 

6 

Mont.: 

Dillon 

24.4 

65.1 

16.67 

7 


Ennis 

21.7 

64.5 

10.69 

8 


Polaris 

20(ca.) 

62(ca.) 

?3 

9 

Wyo.: 

Centennial 

21.5 

61.4 

16.91 

10 

Colo.: 

Boulder Co, 

28(ca.) 

71(ca.) 


11 


Red Feather L., 8400 ft. 

15.0 

57.9 

^4 

12 


Ft. Collins 

26.0 

68.9 

15.20 

13 

Utah: 

Stockton 

25.5(ca.) 

72.3(ca.) 

12.90^ 

14 

N. Mex. 

: Cloudcroft 

30.1 

59.7 

24.58^ 


1, Temperature values are those of Spokane reduced equally by a lapse 
rate of 1° P/300 ft. Precipatation estimated from regional values. 

2, Spots for Loneroek^ Oregon, and Cloudcroft, New Mexico, are omitted 
from the graph. They fall so far outside the pattern of the 
remaining localities that I strongly suspect them of not represent- 
ing the particular localities where the sheridanil were actually 
tidcen. There is no error in the values themselves which were 
kindly confirmed for me by Mr. T. L. Long, National Weather Records 
Center, Asheville, North Carolina, Needless to say, this is a 
common problem in mountainous country where small differences 

in elevation or exposure can exert major changes in the climatic 
picture. 

3, Estimated values. 

4, Temperature values are those of Ft. Collins reduced as described in 
note 1, 

5, All values are interpolated from those of two adjacent bracketing 
localities . 


158 


CLENCH 


/. Res. Lepid. 



Fig. 1. Map of a portion of the range of C. {Callophrys) sheridanii 
showing localities of subspecies neoperplexa (open circles) and subspecies 
neivcomeri (solid circles). Excluded areas (shaded) have one or more of the 
following: (1) mean January temperatures below 14°F or above 32°; (2) 
mean July temperatures below 57° or above 76°; (3) mean annual precipita- 
tion below 10 or above 25 inches. 

The method of correlated responses makes use of a well known 
fact that organisms seldom respond to these or other climatic factors 
singly, but that tolerance to one is usually in some way related to 
the value of the other. This can be seen in the accompanying graph 
of sheridanii localities ( fig. 2 ) , plotted for values of mean January 
and mean July temperatures. When mean July temperature is below, 
for example, 60^ F, then sheridanii will occur in such localities only 
if mean January temperature is somewhere between, roughly, 14° - 
22°; but when mean July temperature exceeds 70° it occurs in 
localities where the mean January temperature is above about 24°. 
Thus for any given July temperature the range of tolerance to January 
temperatures is much less than the range applicable to the species 
over its whole range and conversely. 


mean Jan. 


2(2):in-]60, 196} 


CALLOPHRYS FROM NORTHWEST 


159 



Fig. 2. Graph of known localities of C. {Callophrys ) sheridanii (numbered 
as in table 1 ) according to their mean January and mean July temperatures. 


160 


CLENCH 


/. Res. Lepid. 


Similar plots (mean January against mean July) have been con- 
structed for several other Norther American lycaenids {Callophrys {In- 
cisalia) ] Lycaena thoe\ all of which show patterns generally similar 
to that of sheridanii. In all, for example, the correlation is rectilinear, 
occupying a band of more or less uniform breadth, though this 
breadth, the slope of the band, and its position vary from one species 
to another. In those species with a sufficient number of locality points 
( sheridanii is not one of them ) a further, rather paradoxical, effect is 
noted: the upper limit of temperature tolerance is set by a January 
threshold, the lower limit by a July threshold. In other words, such 
species can extend into warm areas only to a point where increasing 
winter temperatures form a barrier; and can extend into cold 
areas only so far as summer temperatures continue to be sufficiently 
warm. Neither the heat of summer nor the cold of winter seems 
to be relevant. This may well be true of sheridanii but the available 
data are insufficient to demonstrate it. 

There will be, then, within the limits set by the particular factors 
singly^ additional "excluded areas’’ in this two dimensional manifold 
whose July and January means lie outside this band of correlated 
temperature responses. These, when added to the other excluded areas 
( as shown on the map, fig. 1 ) , would still further restrict the territory 
available to the species. This has not been done for sheridanii because 
the number of locality points available is not sufficient to permit a 
reliable determination either of the central regression line or the 
breadth of the band. Inspection, however, shows that much of the 
above-mentioned areas of Montana and South Dakota would thereby 
be excluded. This, parenthetically, suggests that the correlated re- 
sponse to summer and winter mean temperatures is a major impedi- 
ment to the eastward spread of sheridanii. 


REFERENCES 

BARNES, WILLIAM and FOSTER H. BENJAMIN, 1923. Notes and new 
species. Contrib. Nat. Hist. Lejid. N. America^ : 62-96 [Callophrys, 

pp. 64-69]. 

BLACKMORE, E. H. 1927. Check list of the Macrolepidoptera of British 
Columbia, Victoria-. E. H. Ban field, 47 pp. 

BOWMAN, KENNETH 1919. Annotated check-list of the Macrolepidoptera 
of Alberta. Red Deer: Alberta Nat. Hist. Soc., 16 pp. 

CLENCH, HARRY K. 1944. Notes on lycaenid butterflies, a. The genus 
Callophrys in North America. Bull. Mus. Comp. Zool. 94: 217-229. 

1961. Tribe Theclini in Paul R. and Anne H. Ehrlich, How 

to know the butterflies (Dubuque, Iowa: W. C. Brown Co.): 177-220 
[Callophrys {Callophrys), p.p. 209-210], 

LEIGHTON, BEN V. 1946. The butterflies of Washington, Dniv. Washington 
Publ. Biol. 9: 47-63. 


Journal of Research on the Lepidoptera 


2 ( 2 ) : 161 - 169 , 1963 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 

© Copyright 196 } 

EUPHYES DUKESI 

A Review of Knowledge of its Distribution in 
Time and Space and its Habitat 

BRYANT MATHER 

Box 2131 Jackson, Mississippi 

Euphyes DUKESI Lindsey/ has been reported from Alabama, 
Virginia^ Ohio, Michigan, and Louisiana. This paper reports its occur- 
rence in Mississippi and North Carolina and in a new section of Louisi- 
ana, gives notes on its habitat' and on its known distribution in time 
and space. 

The finding of E. dukesi in Mississippi suggested the desirability 
of reviewing available information on all the known occurrences. In 
the course of the review, valuable unpublished information on its occur- 
rence in North Carolina, Virginia, Louisiana, Michigan, and Ohio was 
obtained and is here presented. It is hoped that this review will stimu- 
late search for additional occurrences, especially in the wide gaps that 
separate the known occurrences. 

ALABAMA 

Euphyes dukesi was described by Lindsey (1923) from a type series 
of four specimens from Mobile Co., Alabama, that were taken by W. C. 
Dukes of Mobile. Bell (1926) gave additional data on the type locality 
and reported having taken specimens on Hibiscus flowers one mile 
from Union Station, Mobile, in a marsh with waist- to shoulder-high 
grass. Lindsey, Bell, and Williams ( 1931) reported that it was collected 
in considerable numbers in 1925 by Dr. Van Aller and Mr. Lading 
(sic)^ of Mobile and by Bell during a short collecting trip near that 
city. It was on the wing from 24 August to 11 October. Holland 
(1931) figured a S, referred to in his text (p. 388) as "type” and 
on his plate (plate LIV, fig. 26) as "paratype” and stated: "It occurs 
in southern Alabama and probably elsewhere along the Gulf.” Klots 
(1951) figured a pair from Mobile, Ala. Evans (1955) recorded that 
2 S $ and 1 $ from Alabama were in the British Museum. Clench 
(in litt) reported that the only two specimens of dukesi in the Car- 
negie Museum are the two para types mentioned by Lindsey (1923) 

as being in his collection. 

'This species was formerly placed in the genus Atrytone Scud. 1872. Evans (195 5 noted that 
the type species of Atrytone is arogos B. & LeC. 1833. He included Atrytone only arogos and 
logan. He also noted that the type species of Euphyes Scud. 1872 is vestris Bdv. He included in 
Euphyes the remaining species of North American skippers formerly lumped with arogos and logan 
under Atrytone. The validity of the separation of the species formerly so lumped into the two 
groups into which Etans divided them, and hence the validity of the change of the generic name 
for dukesi from Atrytone to Euphyes appears to be concurred in by all of those with whom the 
author corresponded in collecting the information summarized in this paper. 

2Henry Peter Loding, author of Catalogue of Beetles of Alabama. Ala. Geol. Surv., Monograph 11, 
1945, 172 pp. 


161 


162 


MATHER 


/. Res. Lepid. 


VIRGINIA 

Clark (1932) did not list duke si as possible for the vicinity of 
the District of Columbia. Clark and Clark (1951) described the 
area in Norfolk and Princess Anne Counties from which they had 
dukesi records based on their own work and that of Otto Buchholz. 
They concluded that, in this area, dukesi has one brood that flies 
between 12 June and 16 July. Martin and Truxal ( 1955) reported 
that the Los Angeles County Museum collection included dukesi taken 
in Virginia in August, Rindge checked the American Museum of 
Natural History series and reported (in Litt) that there were 35 S $ 
and 26 $ $ . Virginia specimens from the Buchholz collection include 
both S S and $ $ taken in June for which the localities are given 
as "Norfolk Co.,” "Pr. Anne Co.,” and "Suffolk.” Suffolk, Va., is in 
Nansemond Co. A $ , probably collected by Buchholz, that came to 
us from C. F, dos Passos, is labelled "Pecaty Road, vie. Hickory, Va. 
Norfolk Co., 26 VI 40” Charles V. Covell, Jr. reported (in litt) that 
he had not taken dukesi in his collecting in the Norfolk area. Col. S. S. 
Nicolay reported (in litt) that he had taken specimens in Virginia in 
all months, May through August, is convinced that, in this area, dukesi 
is double brooded, and estimated that the first brood is out between 25 
May and 5 July and the second between 20 August and 15 September. 
He thought that it may be possible for it to be found as late as October. 
Col. Nicolay provided the following notes on collecting dukesi in 
Virginia: "A fair number of specimens were taken in the period 
4-15 July 1951. The location of the habitat is the difficult factor; 
once the habitat is located, they are not particularly difficult to 
capture. North Landing, Princess Anne Co., a locality mentioned by 
Clark and Clark (1951), is actually a bridge crossing a canal. Dukesi 
is found abundantly in the few yards of marsh on either side of the 
bridge and along the banks of a small stream that empties into the 
canal at the bridge, but it is not found 100 yards away from the 
bridge on either side, nor in any of the fields a block away. Dukesi was 
also found in other spots in and around the Dismal Swamp area of 
Princess Anne and Norfolk Counties. Each locality is actually a 
Spot since the insects never wander away from their chosen ground. 
One can collect in a field or along a trail within a stone’s throw of 
the deep swamp habitat of dukesi and not find a single stray. They 
were not found more than a few yards from standing water. The 
best collecting method is to work the edges of small roads that cut 
through the gum swamps. The ^ in search "of 9 9 will fly along 
the edge of the road for a few yards before turning- back into the 
tall swamp grass from whence they came. This grass grows under the 
trees of the swamp, not out in the open. They are fond of pickerel 


2 ( 2 ): 161 - 169 , 1963 


EUPHYES DUKESI 


163 


weed {Pondeteria cor data) and at times were found swarming around 
clumps of its spired blue flower along the edges of small bodies 
of water. The specimens taken in July 1951 were quite well worn, 
especially the $ $ , with an occasional $ appearing fresh. On 25 
August 1951, at the same localities, freshly emerged specimens were 
found in large numbers, particularly $ S . These relations have 
been found to exist in each of the five seasons during which obser- 
vations have been made.” 

OHIO 

References to the occurrence of dukesi in Ohio were found in 
Freeman (1942), Harris (1950), Klots (1951), and Pliske (1958 
”1957”), hut no detailed published account has been located. The 
earliest record of dukesi from Ohio that was found is of a ^ taken 
on 10 July 1940, at Sylvania by Donald Eff given in Freeman ( 1942) , 
Sylvania is 11 miles northwest of Toledo, in Lucas Co., close to the 
Michigan border. Eff described (in litt) the locality as "in the Oak 
Openings in northwest Ohio, an acid, quicksand base, area of about 140 
square miles set in the midst of rich farming land. Before canals 
were dug and the water level was higher, it was a combination of 
swamps and higher sandy knolls. Much of the land in the northeast 
corner of Indiana and southeast Michigan would fall in the same general 
category.” The occurrence of dukesi in Ohio was apparently not 
known to Macy and Shepard (1941). E. S. Badger of Kokomo, 
Ind., described (in litt) his experiences collecting dukesi in Ohio on 
7 July 1957 as "one of the most fantastic experiences I have had in 
this section of the country. Homer Price took me to the old disused 
bed of the Miami Canal on a blistering hot sticky day. Equipped with 
knee-high rubber boots, we followed the canal to an area where it 
was over-grown with wide-bladed sedges from bank to bank. Sloshing 
around in water 6 to 12 inches deep, we started seeing c? flying 
all over the place. The canal was pretty well shaded by trees and 
was only about 12 to 15 feet wide. There were some sedges in the 
adjoining woods in one area and here, particularly, dukesi abounded. 
It had been our intention to take only a half dozen specimens but 
the skipper was so abundant that I ended up with 36 $ $ and 1 $ 
in about 90 minutes of collecting. The only other butterflies were 
common dark skippers, a few worn Strymon falacer, and worn Speyeria 
cybele.” One $ collected by Badger on 7 July 1957, now in our 
collection, is labelled: 'Miami Canal, Paulding Co., Ohio.” Mr. Badger 
not only sent us the foregoing account but also asked Mr. Homer F. 
Price of Payne, Ohio to provide additional information. Mr. Price gave 
(in litt) the following records and comments: 

(a) Marie DeLarme Creek, Carryall Township, Paulding Co., 


164 


MATHER 


}. Res. Lcphl. 


Ohio: 29 June 1955 9 , 4 July 1955 3 6 6 , 30 June 1956 6 , 8 
July 1956 6 , the sedges are nearly shoulder-high some years at this 

station, low ground along the creek. A small colony; in some seasons 

none are seen. 

(b) Miami Canal, Defiance Township, Defiance Co., Ohio: 8 
July 1956 S . Very wet woods along the canal, the sedges were 
rather short, none were seen in the canal, which was dry. 

(c) Miami Canal at Charloe, Brown Township, Paulding Co., 

Ohio: 14 July 1950 c? 9,16 July 1950 2 o o , 23 July 1950 2 5 c? 

30 July 1950 9, 25 August 1950 5 (worn, latest record), 2 July 

1959 3 5 5 19 (fresh). 'The species is quite common some seasons 
in and along the canal where the sedges are of normal size. They 
occur at rather open spots where there is a little sunshine. Many 
specimens have been collected at this station. 

"I am of the opinion that this species is more plentiful in this 
area than is generally believed. Recently I have collected dukesi at 
various stations along the canal from the Defiance County line .south 
to the Putnam County line. This area has a low gradient, having been 
a part of the Black Swamp. The soil is a very heavy clay. I have been 
of the opinion that dukesi probably appeared first in v/estern Ohio 
near the southern terminus of the Miami Canal at Cincinnati and 
extended its range northward along the canal. Hot weather, mud 
and water, mosquitoes and deer flies, probably cause some collectors 
to remain at home during early July when dukesi should be searched 
for. It appears that we have only one brood here. They seem to 
like shade - but not deep shade.” 

Badger gave (in litt) the following additional Charloe records 
from specimens now in his collection that were taken by Price: 15 
July 1951 2 5 5 , 22 July 1951 9, 21 July 1954 9, and 4 July 
1955 5 . Badger has included dukesi on his preliminary manuscript 
list of Indiana butterflies on the basis of its known occurrence in Ohio 
within a few miles of the Indiana border. Price also mentioned that 
he had correspondence with Chermock and had provided material 
for study in connection with the possibility that the Ohio and Alabama 
populations might be subspecifically different. Coveil stated (in litt) 
that his collection contained a 5 obtained from Frank Chermock 
labelled "Chasloe, Pauling Co., Ohio, 15 July 1957” taken by Price. 

LOUISIANA 

Lambremont (1954) described the single speciment known to him 
from Louisiana, taken by him, as constituting a possible westward 
range extension. His specimen^ a 5 , Tulane University #2064, was 
taken IIV2 miles west of Oak Grove, West Carroll Parish, on 19 
June 1950. This locality is near the Boeuf River about 20 miles west 


2 ( 2 ); 161 - 169 , 196 } 


EUPHYES DUKESI 


165 


of the Mississippi River, and more than 200 miles north of the Gulf 
of Mexico. Harris (1950) gave the range of dukesi as: "known 
from southern Louisiana, southern Alabama (Mobile), soudieastern 
Virginia, and Ohio.’’ Harris wrote (in litt) that the information on 
dukesi given in his 1950 report was v/ritten by A. H. Clark and has 
kindly provided a copy of Clark’s letter of 13 July 1950 to him, 
reading in part; "I am returning the manuscript ... I have added 
. . . in brackets ... A. dttkesi which certainly occur (s) in Georgia.’’ 
Harris commented, with regard to Clark’s reference to "southern 
Louisiana” that he assumed that Clark knew of records. It seems un- 
likely that Clark on 13 July 1950 would have known of Lambremont’s 
19 June 1950 record. .The reference by Clark is almost certainly to 
a previously unpublished occurrence discovered by William D. Field 
which Mr. Field has described (in litt) as follows: "I took a series of 
dukesi near a drainage ditch in what had once been a swamp near 
Camp Plauche, Jefferson Parish, La., on May 15, 1944. This swamp 
had been well-drained but had a good flora resembling a woods with 
only a light amount of undergrowth. The details of the latter escape 
me except for the catbrier {Smilax). Camp Plauche was right next to 
the town of Harahan, less than a mile from the Mississippi River. This 
series was sent to Austin Clark for the museum. However, at present 
there is only a single male in the collection.” Harahan is on the 
north (left) bank just a little upstream from New Orleans. 

MICHIGAN 


Pliske (1958 "1957”) described the occurrence of dukesi at Ann 
Arbor, Michigan. He reported 2 S S taken by Arthur Slater and 
himself on 21 July 1956 in a small marsh on the north side of the 
Huron River about one mile east of Ann Arbor. Washtenaw Co., 
Mich.; a second $ taken at the same locality the following day; and 
a $ taken on 13 July 1957. A second locality, Vj mile east of the 
first, yielded a 2 on 27 July 1956 and a $ on 15 July 1957. Both 
marshes were dominated by the lake-margin sedge {Carex lacustris) 
characterized by broad blades, 1 cm. or more in width, spreading in 
a criss-cross manner. On 28 July a 2 was observed ovipositing on 
the underside of a sedge leaf about HA feet above the ground. The 
eggs were collected by W. H. Wagner and were described by Pliske. 
Pliske’s account refers to the Ann Arbor colony as the fourth to have 
been discovered, actually, although not known to him, it was the 
sixth, the fourth and fifth having been the two Louisiana occurrences. 
Wagner has provided (in litt) additional data on the Michigan occur- 
rence of dukesi as follows: Subsequent to the development of the 
data reported by Pliske, a battered 6 that had been taken by Sherman 
Moore on 28 July 1952 at Highland Park, Oakland Co., Michigan 


166 


MATHER 


/. Res. T.epitf. 


was found in the University of Michigan collection in the E dion 
series. Wagner notes that the thoroughly urbanized region of Highland 
Park in Detroit is wholly unsuited to the occurrence of dukesi and 
that the specimen must be a stray. Wagner having visited the North 
Landing, Virginia, locality with Clark and also the Ann Arbor, Mich- 
igan locality, compared them as follows; 'The habitat there (North 
Landing) was very different from the Michigan one: deep, shady 
swamp, the butterflies rather slow flying among grasses and sedges. 
Here the species reminds me more of dion. It flies with the latter 
as well as A. logan, P. viator, E. conspicua, and P. massasoit!’ 

MISSISSIPPI 

Mather and Mather ( 1958) regarded L. dukesi as of probable 
occurrence in Mississippi. This was proved correct when I took a $ 
on 4 October 1959 about 4 miles north of Clinton, Hinds Co. The 
locality is on the north side of the Kickapoo Road bridge over Bogue 
Chitto Creek, Sec. 3.5, T 7 N, R 1 W. The specimen was taken at 
about 3 p.m., feeding on flowers of Eupatorium {Conodinum) coel- 
estinum that were growing in a marshy area within a few feet of a 
pool of essentially stagnant water in the creek bed. This locality is 
about 200 miles north of the Gulf of Mexico. A comparison of the 
three males at hand reveals that the Mississippi specimen is more like 
the Virginia one than the Ohio one especially in that it has less yellow 
hair on the hindwing above and less bright yellow on the hindwing 
below. The separation of the black areas of the stigma on the fore- 
wing above is most distinct in the Virginia specimen, slightly less 
distinct in the Ohio specimen, and still less distinct in the Mississippi 
specimen. At the same locality on the same date, the first Mississippi 
specimen of Poanes viator was taken. A second viator was taken there 
on 11 October 1959 by M. & E. Roshore but no additional specimens of 
either species have been found on numerous subsequent visits. 

NORTH CAROLINA 

Nicolay reported (in litt) the taking of dukesi on 6 June I960 
at a locality about 7 miles southeast of New Bern, Craven Co., in 
a small swamp located along the Neuse River. Based on his observa- 
tions in Virginia he regarded the locality as a typical dukesi habitat. 
Specimens of both sexes were taken, the condition of the $ $ sug- 
gested that they had been on the wing for about a week, the $ $ 
were quite fresh. 


2(2)J6l-169, 196} 


EUPHYES DUKESI 


167 


SUMMARY AND DISCUSSION 

The geographical distribution of the localities at which E. dukesi 
has been found to occur is shown by figure 1; also shown are the wide 
areas between the Great Lakes, the Middle Atlantic Coast, and the Gulf 
States from which the insect is not yet known. Nicolay reported (in 
litt) that he did not find dukesi during the two years (1 947-49 ) 
that he collected in the Pensacola, Fla., area. Zeiger reported (in 
litt) the taking of a series of 17 c? ^ and 5 9 $ of P. viator on 
23 May 1962 among giant cutgrass, in brackish water, along the edge 
of the Suwanee River, near its mouth. He will visit this locality 
again and check for the occurrence of P. dukesi. Neither Lambremont 
(1954) nor Mather and Mather (1958) found it in coastal Louisiana 
or Mississippi. Freeman (1951) did not report it from Texas. Harris 
reported (in litt) that he felt certain that it will be found in swamps 
in Georgia such as those of the Altamaha River, the Okefenokee, 
and other suitable localities. He also indicated that Buchholz, whose 
records of dukesi from Virginia were cited by Clark and Clark (1951) 
and who had many Georgia records, apparently had no records of 
dukesi from Georgia. Roever (in litt) reported that he had never 
collected in the type of marsh in Tennessee in which dukesi might 
be expected to occur but regarded the open marsh around Reelfoot 
Lake in northwest Tennessee and the spring-fed gum swamps in 
the Hatchie River bottom near Viedo, 4-5 miles southeast of Mercer, 
Madison Co., as likely areas. 

E. dukesi appears to be single brooded in Ohio and Michigan and 
double-brooded in Virginia and the Gulf states; the months of record 
for the seven states in which it is known to occur are shown in table 1. 

E dukesi appears to be a swamp butterfly, but while the Ohio 
and Michigan occurrences are apparently closely connected with sedges, 
the Virginia occurrence is in the gum (Nyssa aquatica) swamps, which 
as Clark and Clark (1951, p. 8) pointed out, lie west of the sedge 
marshes that, in turn lie behind the sand dunes of the Virginia coast. 
Until more complete knowledge of the distribution of dukesi becomes 
available, it would seem prudent to describe its distribution as: north- 
east and southeast Louisiana, central Mississippi, southwestern Ala- 
bama; eastern North Carolina, southeastern Virginia; northwestern 
Ohio, and southeastern Michigan, rather than in terms that imply its 
occurrence in the large regions between these areas of known occur- 
rence. 


168 


MATHER 


J. Res. LepiiJ. 



Fig. 1. Map showing the locations where Euphyes dukesi 
has been found. 



MAY 

JUN 

JUL 

AUG 

SEP 

OCT 

MICHIGAN 



X 




OHIO 


X 

X 

X 



VIRGINIA 

X 

X 

X 

X 

(a) 

(b) 

NORTH CAROLINA 


X 





ALABAMA 




X 

X 

X 

MISSISSIPPI 






X 

LOUISIANA 

X 

X 






(a) Probable (b) Possible 


TABLE 1. The monthly occurrence of Euphyes dukesi at 
various localities in the eastern United States. 


2(2): 161-169, I96.i 


EUPHYES DUKESI 


169 


ACKNOWLEDGEMENTS 

The principal value of this paper derives from the data provided 
by Col. S. S. Nicolay, Cherry Point, N. C.; Mr. William D. Field, 
Washington, D. C.; Mr. F. S. Badger, Kokomo, Ind.; Mr. Homer F. 
Price, Payne, Ohio; Mr. Lucien Harris, Jr., Atlanta, Ga.; Mr. Charles 
V. Coveil, Jr., Norfolk, Va.; Dr. Frederick H. Rindge, New York, 
N. Y.; Mr. Charles F. Zeiger, Jacksonville, Florida; Mr. Harry K. 
Clench, Pittsburg, Penn.; Mr. Kilian Roever, Tucson, Ariz.; Dr. 
Warren H. Wagner, Ann Arbor, Mich; Mr. Donald Eff, Boulder, 
Colo.; Mr. H. A. Freeman, Garland, Texas; Mr. John M. Burns, Middle- 
town, Conn.; and their kind permission to publish them; their coopera- 
tion is deeply appreciated. 


REFERENCES 

BELL, E. L. 1926. Notes of some Hesperiidae from Alabama, Jour. N. Y. 
EnL Soc., 34: 269-271. 

CLARK, AUSTIN H. 1932. The Butterflies of the District of Columbia and 
vicinity. Smithson. Inst., Bull. U. S. Nat. Mus., 157, 337 pp. 

CLARK, AUSTIN H. and LEILA F. CLARK 1951. The butterflies of Vir- 
ginia. Smithson. Misc. Coll., 116 (7): 1-239. 

EVANS, W. H. 1955. A Catalogue of the American Hesperiidae, Part IV. 
British Museum- (London) , 499 pp. 

FREEMAN, H. A. 1942. Notes on some North American Hesperiidae with 
the description of a new race of Polites verna ( Edwards ) ( Lepidoptera ; 

(Rhopalocera) . Ent. News, 53: 103-106. 

_ — 1951. Ecological and sytsematic study of the Hesperiidea or 

Texas. Sotithern Methodist Univ. Studies, No. 6: 1-67. 

HARRIS, LUCIEN JR. 1950. The butterflies of Georgia. Bull. Georgia Soc. 
Naturalists, 5 : 29 pp. 

HOLLAND, WILLIAM J. 1931. The Butterfly Book (revised edition). 

Douhleday & Co., Garden City, N. Y., 424 pp. 

KLOTS, ALEXANDER B. 1951. A Field Guide to the Butterflies. Houghton 
Mifflin Co., Boston, 349 pp. 

LAMBREMONT, EDWARD N. 1954. The butterflies and skippers of Louis- 
iana. Tulane Stud. ZooL, 1: 125-164. 

LINDSEY, A. W. 1923. New North American Hesperiidae. Ent. News. 
34: 209-210. 

ERNEST L. BELL and R. C. WILLIAMS, JR. 1931. The Hes- 

perioidea of North America. Jour. Sci. Lab. Denison Univ. 26: 1-142. 
MACY, RALPH W. and HAROLD H. SHEPARD 1941. Butterflies. Univ. of 
Minnesota Press, Minneapolis, 247 pp. 

MARTIN, LLOYD M. and FRED S. TRUXAL 1955. A list of North Amer- 
ican Lepidoptera in the Los Angeles County Museum, Part 1 - Butterflies. 
Los Angeles Museum, Science Series, No. 18: 1-35. 

MATHER, BRYANT and KATHARINE MATHER 1958. The butterflies of 
Mississippi. Tulane Stud. Zool. 6: 63-109. 

PLISKE, THOMAS E. 1958 ”1957”. Notes on Atrytone dukesii (sic), a rare 
species new to southern Michigan (Hesperiidae). Lepid. News, 11: 42. 


Journal of Research on the Lepidoptera 


2 ( 2 ) : 170 - 172 , 1963 


1140 W. Orange Grove Ave., Arcadia, California, U.S.A. 
© Copyright 1 ^ 6 } 


THE COMPLETE LIFE HISTORY OF STAPHYLUS 
HAYHURSTI 


JOHN RICHARD HEITZMAN 

3112 Harris Ave., Independence, Missouri 


This obscure little skipper is one of those species that are 
easily overlooked by the non-observant collector. It inhabits a wide 
range in the eastern United States extending at least as far as Pennsyl- 
vania and Nebraska and south to Florida and Texas, being replaced 
in southern Texas by the closely related Staphylus mazans Reakirt. Host 
plants include Chenop odium and Alternanthera and the species is rather 
localized to areas where these weeds grow. One of the peculiarities of 
hayhursti is the moth like habit of spreading the wings out flat against 
the surface upon which it alights. It is strongly attracted to certain 
flowers and at times I have found a dozen or more specimens clustered 
about a bed of Spearmint flowers {Mentha spicata) during late July 
in a good local. Sweet Clover (Melilotus) also seems to have a par- 
ticular attraction for the species. In rural areas the best collecting 
spots for hayhursti will be associated with moist rather shady habitat. 
An idea spot is a small creek running through a wooded area or the 
weedy edges beside an old railroad track. In urban areas it is drawn 
to vacant lots grown up in weeds where an ample supply of the host 
plants can be found growing. There are two distinct broods produced 
each year in Missouri and probably at least three further south. In a 
normal year you can find hayhursti from the third week in May to 
the end of the third week in June and again from mid July until the 
end of August. The larvae produced by the second brood normally 
hibernate in the third instar in this area. An occasional third brood 
specimen is produced but these are very rare. 

From time to time scattered bits of information on the early stages 
of this species have appeared but as far as I can ascertain there has 
be